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cryptogamie
mycologie
volume 36 n°1 2015
contents
Asanka R. BANDARA, Sylvie RAPIOR, Darbhe J. BHAT, Pattana
KAKUMYAN, Sunita CHAMYUANG, Jianchu XU & Kevin D. HYDE —
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom with
Multiple Developed Health-Care Products as Food, Medicine and
Cosmetics:areview........................................3-42
Marisa de CAMPOS-SANTANA, Gerardo ROBLEDO, Cony DECOCK &
Rosa Mara BORGES DA SILVEIRA — Diversity of the poroid
Hymenochaetaceae (Basidiomycota) from the Atlantic Forest and Pampa in
SouthernBrazil...........................................43-78
Rosa Emilia PÉREZ-PÉREZ, Gonzalo CASTILLO-CAMPOS & Marcela
Eugenia da Silva CÁCERES — Diversity of corticolous lichens in cloud
forest remnants in La Cortadura, Coatepec, Veracruz, México in relation to
phorophytes and habitat fragmentation. . . . . . . . . . . . . . . . . . . . . . . . . .79-92
Wen-Jing LI, Sajeewa S. N. MAHARACHCHIKUMBURA, Qi-Rui LI,
D. Jayarama BHAT, Erio CAMPORESI, Qing TIAN, Indunil C.
SENANAYAKE, Dong-Qing DAI, Putarak CHOMNUNTI & Kevin D. HYDE
— Epitypification of Broomella vitalbae and introduction of a novel species
of Hyalotiella............................................ 93-108
Michel Navarro BENATTI, Márcia Isabel KÄFFER, Suzana Maria de
Azevedo MARTINS, Alessandra Bittencourt de LEMOS — Bulbothrix
bulbillosa, a presumed Galapagos endemic, is common in Rio Grande do
Sul State, Brazil (Parmeliaceae, lichenized Ascomycota). . . . . . . . . . . . . .109-114
Instructions aux auteurs / Instructions to authors. . . . . . . . . . . . . . . . . . . .115-118
Cryptogamie, Mycologie, 2015, 36 (1): 3-42
© 2015 Adac. Tous droits réservés
doi/10.7872/crym.v36.iss1.2015.3
Polyporus umbellatus,an Edible-Medicinal Cultivated
Mushroom with Multiple Developed Health-Care
Products as Food, Medicine and Cosmetics: a review
Asanka R. BANDARAa,b,c, Sylvie RAPIORd, Darbhe J. BHATe,
Pattana KAKUMYANa, Sunita CHAMYUANGa,
Jianchu XUb,c & Kevin D. HYDEa,b,c*
aInstitute of Excellence in Fungal Research, Mae Fah Luang University,
Chiang Rai 57100, Thailand
bWorld Agroforestry Center, East and Central Asia Region, Kunming 650201,
Yunnan, China
cCenter for Mountain Ecosystem Studies, Kunming Institute of Botany,
Chinese Academy of Science, Kunming 650201, Yunnan, China
dCEFE UMR 5175, CNRS – Université de Montpellier –
Université Paul-Valéry Montpellier – EPHE, Laboratoire de Botanique,
Phytochimie et Mycologie, Faculté de Pharmacie, BP 14491,
15, avenue Charles Flahault, F-34093 Montpellier Cedex 5, France
eFormerly, Department of Botany, Goa University, Goa 403 206, India;
No. 128/1-J, Azad Housing Society, Curca, P.O. Goa Velha 403108, India
Abstract –Polyporus umbellatus is a medicinal mushroom belonging to the family
Polyporaceae which forms characteristic underground sclerotia. These sclerotia have been
used in Traditional Chinese Medicine for centuries and are used to treat edema and
promote diuretic processes. Over the past few decades, researchers have found this taxon to
contain many bioactive compounds shown to be responsible for antitumor, anticancer,
antioxidant, free radical scavenging, immune system enhancement and antimicrobial
activities. Due to its promising medicinal value, P. umbellatus is used as an ingredient in
many medicinal products and food supplements. Thus demand for P. umbellatus has
increased. To supply the high global demand, P. umbellatus is cultivated under natural or
industrial conditions. In this review we discuss optimal conditions for the cultivation and
culture of P. umbellatus. We also focus on the medicinal uses of P. umbellatus, the diversity
of bioactive metabolites with various pharmacological properties and the medicinal
products of great interest for health care or as alternative drugs.
Anticancer / Antimicrobial / Antioxidant / Antitumor / Bioactive molecules / Immunity /
Medicinal mushroom / Polysaccharides / Steroid
* Corresponding author: email addresses: Kevin D. HYDE, kdhyde3@gmail.com
4A. R. Bandara et al.
INTRODUCTION
Mushrooms are defined as macrofungi with distinctive surface or
subterranean fruiting bodies, large enough to be seen with the unaided eye. There
are an estimated 140,000 species (Chang & Miles, 1992; Hawksworth, 2001;
Wasser, 2002), but about 10% are thought to be named (Hawksworth, 2001;
Lindequist et al., 2005; Wasser & Didukh, 2005). The total number of edible and
medicinal species is over 2300 Ça∑larirmak, 2011; Ying et al., 1987; Maass et al.,
2012). Mushrooms provide dietary protein, essential amino acids, carbohydrates,
vitamins and minerals (Ça∑larirmak, 2011; Cheung, 2008; Jong & Birmingham,
1990; Luangharn et al., 2014; Smith et al., 2002; Thatoi & Singdevsachan, 2014).
Several thousands of years ago people in the Orient recognized that many edible
and certain non-edible mushrooms have valuable health benefits (Cao et al., 2012;
Hobbs, 1995; Mortimer et al., 2012; Smith et al., 2002; Thawthong et al., 2014;
Alves et al., 2012; Giavasis, 2014; Quang et al., 2006).
Polyporus umbellatus (Pers.) Fr. is a medicinal mushroom in the family
Polyporaceae of class Basidiomycetes (Choi et al., 2003; Ying et al., 1987; Zhou &
Guo, 2009). It is a saprobe(Sekiya et al., 2005; Sun & Yasukawa, 2008) which
causes white rot of wood (Choi et al., 2002; Lee et al., 2007; Lee et al., 2005;
Ryvarden & Gilbertson, 1994; Zhao & Zhang, 1992). Scopoli (1772) initially
named this fungus as Boletus ramosissimus. It was renamed as Fungus
ramosissimus by Paulet (1793), Boletus ramosus by Vahl. (1797), and Boletus
umbellatus by Persoon (1801, 821). Finally it was named Polyporus umbellatus by
Fries, and this name has since been used (Partnership; Murrill, 1904; Robert et al.,
2005).
Polyporus umbellatus is commonly referred to as Grifola umbellata
(Hall et al., 2003; Roody, 2003; Xing et al., 2012) and more infrequently
Dendropolyporus umbellatus (Pouchus, 2012; Roody, 2003; Xing et al., 2012). Its
common names include Zhuling ()(HogTuber) in Chinese, Chorei-Maitake
(wild boar dung Maitake) or Tsuchi-maitake (Earth Maitake) in Japanese,
(Miyazaki & Oikawa, 1973; Stamets, 2000; Stamets, 2002) Eichhase in German,
(Bachmeier et al., 2011) Polypore en ombelle in French (Pouchus, 2012) and
umbrella polypore in English (Fischer & Bessette, 1992; Lincoff, 2010; Lincoff &
Nehring, 2011).
Polyporus umbellatus is widely distributed in the temperate regions of
the Northern hemisphere in Asia, Europe and North America (Stamets, 2000;
Zhao et al., 2009c; Zhao et al., 2009e; Zhou et al., 2007; Kikuchi & Yamaji, 2010).
In Asia, it has been recorded in China, (Ying et al., 1987; Zhang et al., 2010b;
Zheng et al., 2004; Zhou & Guo, 2009) India, (Núñez & Ryvarden, 1995; Núñez
& Ryvarden, 2001) Japan, Korea and USSR (Kikuchi & Yamaji, 2010; Zhao &
Zhang, 1992). In Europe it has been recorded in southern and central Europe,
northern to most southern coastal areas of Fennoscandia, (Ryvarden &
Gilbertson, 1994) Poland, (Zhao & Zhang, 1992) France, the UK (Courtecuisse,
1999) and Slovakia (Kunca, 2011). It has been found in north central and north-
eastern parts of North America (Ryvarden & Gilbertson, 1994) and recorded
from east Canada to Tennessee (Lincoff & Nehring, 2011). Gilbertson and
Ryvarden reported this mushroom in Montana and Washington State (Stamets,
2000). The fungus has also been reported from Ithaca, (Murrill, 1904) Amherst,
(Hitchcock, 1829) Ohio, Iowa, Idaho in USA (Lincoff & Nehring, 2011).
Polyporus umbellatus prefers relatively warm regions in broad-leaved (Jong &
Birmingham, 1990; Kikuchi & Yamaji, 2010; Kunca, 2011) and coniferous forest
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom: a review 5
(Stamets, 2000; Ying et al., 1987). Polyporus umbellatus has been observed in
deciduous forests, (Stamets, 2000) as sclerotium close to the stumps of hardwoods
such as Alnus, Carpinus, Castanea, Fagus and Quercus.Quercus seems to be the
favorite host of P. umbellatus and on a few occasions it has also been detected
under Picea and Pinus trees (Núñez & Ryvarden, 1995; Overholts, 1914;
Ryvarden & Gilbertson, 1994; Zhao & Zhang, 1992). The fungus has also been
found around the roots of alder and Japanese oak (Ueno et al., 1980). It prefers
dead roots or buried wood, and birch, maples, willow and beech stumps (Stamets,
2000; Ying et al., 1987). The fungal fruiting portion found on the ground is edible,
whilst the underground part has medicinal properties (Jong & Birmingham, 1990;
Ying et al., 1987).The relative incidence of this fungus was higher in hilly terrains
than in lowlands, while it was found only rarely in the uplands of Slovakia (Kunca,
2011). Polyporus umbellatus has been found in soil rich in lignicolous organic
matter (Stamets, 2000) and mostly in acidic soils (Kunca, 2011). Through a
diversity study carried out on P. umbellatus strains obtained from various parts of
China, it was established that the fungus exhibits an uneven and high genetic
diversity and an abundant environmental heterogeneity (Xing et al., 2013a; Zhang
et al., 2012b). Molecular analysis based on rDNA data indicates significant inter
and intra population variation. The nucleotide diversity was usually higher in the
ITS sequences than in the 28S rDNA sequences (Xing et al., 2013a).
Fossil records reveal that Polypores existed 300 million years ago, during
the carboniferous age when the evolution of woody gymnosperms began (Zhao &
Zhang, 1992). The genus Polyporus has been categorized into five groups and
P. umbellatus belongs to a group which has branched stipes and/or sclerotia. Only
P. umbellatus and P. mylittae form basidiocarps on sclerotia. Spore size, and the
presence of a white to brownish pileus covered with fine scales distinguish
P. umbellatus from P. mylittae (Zhao & Zhang, 1992). Macroscopically, Grifola
frondosa (Maitake) appears to be a close relative of P. umbellatus, (Murrill, 1904)
but biologically the two have different life cycles (Stamets, 2000).
REVIEW
The fungus
Mycelium
Polyporus umbellatus produces white, longitudinally linear mycelia on
agar, in grain, and in sawdust media that soon become densely cottony, thick and
peelable. When maturing on sterilized sawdust, concentric rings appear with an
outer layer of yellowish, gelatinous exudate (Guo et al., 2002; Stamets, 2000). It
has a musty, sour, slightly bitter, and unpleasant odor (Stamets, 2000). Polyporus
umbellatus mycelium requires a relatively long time to grow in artificial media
(long lag phase) (Huang & Liu, 2007; Wang et al., 2004). The relatively long time
taken for the fusion of monokaryotic hyphae and slow growth of dikaryotic
mycelium are the reasons for this (Xing & Guo, 2008). After this long lag phase,
the conidia (asexual spores) are released by breaking up of the hyphae and a rapid
growth of the mycelium can then be seen (Huang & Liu, 2007; Wang et al., 2004).
Xing and Guo revealed that the conidia of P. umbellatus could be produced from
dikaryotic mycelia (Xing & Guo, 2008). Huang and Liu discovered an artificial
medium which possesses the ability to shorten the lag phase in both the solid-state
6A. R. Bandara et al.
and submerged culture of P. umbellatus (Huang & Liu, 2007). Zhang et al.
reported the production of asexual spores and calcium oxalate crystals from the
mycelium on potato dextrose agar (PDA) (Zhang et al., 2010b). Polyporus
umbellatus mycelia growing in PDA excrete intra and extracellular
polysaccharides (Zhang et al., 2010b). Correlation analysis indicate that intra and
extracellular polysaccharide content had significant and positive relationship, but
extracellular content was negatively correlated with daily mycelial growth rate
(Zhang et al., 2010b).
Sclerotia
Polyporus umbellatus forms an irregular tuber-like underground
structure known as a sclerotium (Ying et al., 1987; Choi et al., 2002; Choi et al.,
2003). It is one of two Polyporus species which develops fruiting bodies on
underground-buried sclerotia. Similar to other species, the sclerotia of
P. umbellatus are connected with the rotten wood in the ground (Zhao & Zhang,
1992). The sclerotium of P. umbellatus is also formed adhering to the living roots
of deciduous species belonging to genera Quercus and Alnus (Kikuchi & Yamaji,
2010).
The sclerotium of P. umbellatus is an irregular, bumpy, rugged and multi-
branched tuber, woody in texture, the upper surface of which is dark brown to
black and the inner part white (Choi et al., 2002; Imazeki & Hong-
o, 1989; Kunca,
2011; Lee et al., 2005; Ying et al., 1987). The sclerotium of P. umbellatus is mild,
sweet and bland in flavor and when dried is used as a crude drug for medicinal
purposes in the Orient, i.e. China, Korea and Japan (Choi et al., 2002; Jong &
Birmingham, 1990; Kikuchi & Yamaji, 2010; Liu & Liu, 2009; Wu, 2005; Ying
et al., 1987). Sclerotia are formed underground; typically between 10 and 15 cm
deep and are rarely found below a depth of 30 cm (Kikuchi & Yamaji, 2010).
Sclerotia of P. umbellatus comprise hyphae and these hyphae are highly
differentiated in structure. The hyphae are organized and form several distinctive
layers inside the sclerotia (Guo & Xu, 1991). As in other higher fungi, the
development of the sclerotium of P. umbellatus has three distinguishable stages
(Choi et al., 2002). During development, the colour in the sclerotium changes and
a new primordium is formed. Large prismatic crystal structures and thick-walled
cells in the centre of hyphae are formed in contrast with other fungal sclerotia
(Choi et al., 2002; Guo & Xu, 1992a). Sclerotia of P. umbellatus and the forest
pathogenic fungus Armillaria mellea form a symbiotic relationship by means of
mutual assimilation (Guo & Xu, 1992b) and growth of sclerotia depends on this
relationship (Xing et al., 2012). Previous studies have revealed that A. mellea
appeared to be a nutritional factor which improves the growth of the sclerotium
of P. umbellatus (Choi et al., 2002; Guo et al., 2002). Armillaria mellea increases
mycelial growth and the production of metabolites such as ergone (Lee et al.,
2007). With promotion of mycelium growth of P. umbellatus by water extract of
A. mellea rhizomorphs, it has been shown that A. mellea acts as a good carbon and
nitrogen source upon which the growth of P. umbellatus depends (Guo et al.,
2011). When the rhizomorphs of A. mellea are introduced into the sclerotia of
P. umbellatus, the sclerotium forms an enclosed cavity around them, in order to
prevent excess colonization by A. mellea. Furthermore, the rhizomorphs inside the
cavity are degraded and the resultant nutrients are absorbed by the sclerotium
(Xing et al., 2012).
Guo et al. classified another companion fungus (Grifola sp.) associated
with sclerotia of wild P. umbellatus, which is related to sclerotial formation (Guo
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom: a review 7
et al., 2002). This companion fungus induces activation of P. umbellatus enzymes
used in sclerotial formation and differentiation by supplying the nutrient
supplements (Xing & Guo, 2004). Kikuchi and Yamaji discovered that not only
A. mellea, but also five other Armillaria species may have a symbiotic relationship
with P. umbellatus (Kikuchi & Yamaji, 2010). These Armillaria species are
believed to have co-evolved with P. umbellatus and their population structure
selected by nature under specific microenvironments (Zhang et al., 2012b). Feng
et al. also recorded A. mellea and A. gallica associated with sclerotia of
P. umbellatus (Feng et al., 2012). In addition, fungal species belonging to genera
such as Eurotium, Fusarium, Geomyces, Mucor and Penicillium were identified
residing with sclerotia of P. umbellatus, and these fungal communities varied with
host location where observed in China (Xing et al., 2012).
The sclerotia of P. umbellatus can survive in soils for a long time and
have the ability to produce new sclerotia directly from the existing ones under
appropriate conditions (Xiaoke & Shunxing, 2005). After months of dormancy the
sclerotium become soft and swollen and on absorbing water produce fruiting
bodies (Stamets, 2000). Unlike other sclerotium forming fungi, the sclerotia of
P. umbellatus can be reproduced only from sclerotia and not hyphae (Wang et al.,
2004). The sclerotium of P. umbellatus is produced mainly in the provinces of
Shanxi, Henan, Hebei, Sichuan, and Yunnan in China. In the process, the fruiting
bodies collected in the spring and autumn are cleaned, dried, sliced, and used
unprepared (Wu, 2005).
Fruiting bodies
The sclerotium of P. umbellatus swells with water and produces
numerous multi-branched, circular mushrooms with umbellate caps (pilei) (Ying
et al., 1987; Stamets, 2000). It fruits annually (Zhao & Zhang, 1992). The pileus is
fleshy and smooth when fresh, hard and brittle-wrinkled when dry (Núñez &
Ryvarden, 1995; Overholts, 1914; Ryvarden & Gilbertson, 1994; Ying et al., 1987).
The fruiting body is one of the most fragile and delicate of mushrooms of species
in the genus Polyporus (Stamets, 2000). The mushroom is centrally stipitate and
the central part of the cap is concave or subfunnel-shaped (Murrill, 1904; Zhao &
Zhang, 1992). Bouquets of mushrooms arise from a common stem base (Ryvarden
& Gilbertson, 1994; Stamets, 2000). The multiple circular pilei arising from a
common stem make this a very distinct species (Ryvarden, 2014). The stipe is
thick at the base, thinner towards the pilei and richly branched (Núñez &
Ryvarden, 1995). The fruit bodies are whitish at first, becoming brown with age,
with an under side featuring circular to angular pores (Ryvarden & Gilbertson,
1994; Stamets, 2000). Pore surface on drying become brownish to brown; pores
are suborbicular, angular, or irregulary lacerate (Zhao & Zhang, 1992). They are
often waterlogged due to a high water carrying capacity (Stamets, 2000). The
hyphal system of P. umbellatus is dimitic, non-septate and thin or slightly thick-
walled and clamp connections can be observed on the hyaline generative hyphae
(Dai et al., 2014; Ryvarden & Gilbertson, 1994; Stamets, 2000; Zhao & Zhang,
1992). Few gloeoplerous hyphae are also present (Núñez & Ryvarden, 1995).
A study carried out on the fruit body development of P. umbellatus, Guo et al.
concluded that the fruit body possesses all three types of hyphae, known as
trimitic hyphal system (Guo et al., 1998). The fruit body bears clavate-shaped
basidia, with 2-4-sterigmata, and basal clamps, in which basidiospores of
cylindrical, hyaline, thin-walled, smooth and white in deposit are located
(Overholts, 1914; Ryvarden & Gilbertson, 1994; Núñez & Ryvarden, 1995;
8A. R. Bandara et al.
Stamets, 2000). Young fruiting bodies of P. umbellatus are edible (Ying et al.,
1987; Jong & Birmingham, 1990; Zhao & Zhang, 1992). The protein content is
higher than the polysaccharide content in the fruiting body, therefore it has lower
polysaccharide/protein ratio compared with Ganoderma lucidum, Lentinula
edodes, Macrocybe lobayensis, Schizophyllum commune, Trametes versicolor,
Tremella fuciformis and Volvariella volvacea (Liu et al., 1997). The sporocarp
production of P. umbellatus follows that of typical forest macrofungi. The
sporocarp production of P. umbellatus increases significantly during some seasons
and corresponds with weather patterns (Kunca, 2011).
Chemical composition
Fruiting bodies, sclerotium and mycelium of P. umbellatus contain
important bioactive substances which are of different chemical composition and
mode of action. Preliminary studies showed that the P. umbellatus contains 46.6%
coarse fiber, 7.89% coarse protein, 6.64% ash and 0.5% of carbohydrate (Ying
et al., 1987). The sclerotium was investigated for its chemical content; Chen and
Deng reported amino acids, water, crude proteins, fats, fiber and mineral
compounds (Chen & Deng, 2003), while Lee et al. detected 78.2%
polysaccharides, 16.8% proteins and 4% ash (Lee et al., 2004). Others studies have
demonstrated that the major chemical constituents of the P. umbellatus are
polysaccharides and steroids (Zhao et al., 2009f; Zhao et al., 2009c; Zhao et al.,
2010a).
Guo et al. observed the pattern of changing nutrients contents through
the development of cultured and wild sclerotia, and stated that with the increment
of time, sugar and protein contents decrease (Guo et al., 1992). In the first year of
growth the amount of fat and in the consecutive year the polysaccharide content
reach their maximum values. Although the amount of ergosterol is the highest in
the subsequent year in cultured sclerotia, it is lowest in wild sclerotia (Guo et al.,
1992). The first chemical study on P. umbellatus recorded a fatty acid,
2-hydroxytetracosanoic acid [CH3(CH2)21CHOHCOOH], isolated from the
fruiting body of the P. umbellatus (Yosioka &Yamamoto, 1964). Awater-soluble
polysaccharide, a glucan which processed (1→3), (1→4), (1→6)-glycosidic
linkages and branched at C-3 or C-6 positions of glucose residue, was isolated
from the sclerotium of P. umbellatus (Miyazaki & Oikawa, 1973). Kato et al.
obtained D-glucose and small quantities of D-galactose and D-mannose from an
aqueous extract of the sclerotium (Kato et al., 1978). The backbone of these
polysaccharides comprised a β(1→3)linked D-glucose and the authors found
similar β(1→4), β(1→6) linkages which previously recorded (Miyazaki & Oikawa,
1973). Gas chromatography and mass spectrometry analyses revealed that
the polysaccharides of P. umbellatus consist of D-mannose, D-galactose, and
D-glucose at the ratio 20:4:1 (Zhu, 1988). Ohno et al. determined the fungal
(1→3)-β-D-glucan in several edible fungi, including Grifola frondosa and
P. umbellatus, which possess two kinds of conformation in the solid state: helix
(curdlan type) and native (laminaran type) (Ohno et al., 1988; Ohno et al., 1986).
Their findings suggested that the (1→3)-β-D-glucan is the native form in
the fruiting body (Jong & Birmingham, 1990). A patent has been obtained for the
extraction method of β-glucan from the fruiting body of P. umbellatus (Lee &
Park, 2001). Polysaccharides are major components existing in both plants and
animals (Peng et al., 2012) and glucans are one of the major polysaccharide
constituents in the cell walls of fungi (Jong & Birmingham, 1990; Du et al., 2014).
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom: a review 9
From the observation of spectral data, it was concluded that the water-soluble
polysaccharides isolated from the mycelium and sclerotium were similar (Xu et al.,
2004; Tian et al., 2005). Recently, Dai et al. investigated similar polysaccharides in
an aqueous extract of the fruiting body with a molecular mass of 2.27 ×103kDa
containing > 90% D-glucose as its monosaccharide constituent (Dai et al., 2012).
The polysaccharides consist of (1→6, 1→4)-linked-D-glucopyranosyl backbone,
substituted at the O-3 position of (1→6)-linked-D-glucopyranosyl by (1→3)-
linked-d-glucopyranosyl branches and approximately 2930 repeating units; each
containing a side chain of no more than three residues in length (Dai et al., 2012).
Bi et al. isolated P. umbellatus polysaccharides using hot water extracts. According
to Phenol-Sulfuric assay, two compounds were enclosed in this mixture, and they
were placed as GUMP-1-1 and GUMP-1-2 while GUMP-1-1 was comprised of
glucose, mannose and fructose, GUMP-1-2 also included uronic acid and protein
(Bi et al., 2013). Polysaccharide which aqueously extracted from fermented
mycelium and fruiting body of P. umbellatus both consist of glucose and galactose
(Sun & Zhou, 2014). The molecular weight of polysaccharide of mycelium was
857 kDa and molar ratio of glucose to galactose is 1.57:1, while from the fruiting
body, the molecular weight was 679 kDa and molar ratio of glucose to galactose
5.42:1 (Sun & Zhou, 2014).
A method of extracting polysaccharides from the P. umbellatus mycelium
by fermentation in a medium containing soya bean and additives was introduced
by Xu and Zhou (Xu & Zhou, 2003). Similarly, a method of purifying
P. umbellatus polysaccharides using a macroporous resin was discovered (Cui
et al., 2005). A method of extracting highly purified water soluble polysaccharides
was introduced by Wang et al. (Wang et al., 2006). Chen et al. introduced an
ultrasonic extracting technique for polysaccharides of P. umbellatus (Chen et al.,
2008). The polysaccharide content that was extracted by this method was greatly
improved compared with the boiling water method of extraction. Reduced
extraction time, reduced ratio of material to liquid and lowered operating
temperature are other advantages of this method.
Chen et al. discovered polyethylene glycol exhibiting effective stimulatory
effects in mycelial biomass and exopolysaccharides production in submerged
cultures of P. umbellatus (Chen et al., 2010b). Wang et al. recommended optimum
alcohol concentrations, pH values for polysaccharides extraction and fractional
precipitation, (Wang et al., 2010) while Li introduced a technique for extracting
polysaccharides (Li, 2011). Zhang et al. obtained a higher polysaccharide yield
from P. umbellatus using a microwave chemical extraction process (Zhang et al.,
2012a). Quantitative analysis of polysaccharides produced by fermentation of
P. umbellatus mycelium was carried out using HPLC technique (Zhou et al.,
2001). A quantitative analysis of polysaccharides using phenol and sulfuric acid in
P. umbellatus was introduced by Guangwen et al. (Guangwen et al., 2007). This
method is highly sensitive, simple, reproducible and accurate with stable data
(Guangwen et al., 2007). An optimum composition medium which included
Soya bean, used for the production of maximum P. umbellatus mycelium, was
introduced by Zhang and Yu (Zhang & Yu, 2008). Shen et al. found a chemical
treatment to decolorize the precipitated water-soluble polysaccharides from
P. umbellatus mycelium (Shen et al., 2009). Du et al. patented a liquid suspension
culture, which can produce a higher amount of polysaccharides and steroids from
mycelium of P. umbellatus in a short fermentation period (Du et al., 2011).
Steroids are one of the main components of P. umbellatus. Abe et al.
reported that the fruiting body contains ergosta-4,6,8(14),22-tetraen-3-one
(ergone) (Abe et al., 1981). Later, the same compound was isolated from the
10 A. R. Bandara et al.
sclerotia of P. umbellatus (Lee et al., 2005). Ergone is a fungal metabolite derived
from ergosterol (Lee et al., 2005; Lee et al., 2007). Ergone isolated from
P. umbellatus possesses a variety of pharmacological activities, both in vivo and in
vitro, including cytotoxic, diuretic, and immunosuppressive effects (Sun et al.,
2013; Zhao et al., 2011b). Lu et al. isolated four components, viz. ergosta-5,7,22-
trien-3-ol (ergosterol), ergosta-7,22-dien-3-one, ergosta-7,22-dien-3-ol, and 5α,8α-
epidioxyergosta-6,22-dien-3-ol, from the fruiting body of P. umbellatus. Three
of these components (ergosta-7,22-dien-3-ol, ergosta-5,7,22-trien-3-ol, 5α,8α-
epidioxyergosta-6,22-dien-3-ol) were shown to enhance of aggregation of platelets
in rabbits induced by collagen and/or adenosine-5’-diphosphate in vitro (Lu et al.,
1985). The concentration at which the platelets are aggregated by these three
active compounds is less than the effective concentration of cholesterol, which has
a chemical structure very similar to ergosterol.
Ohsawa et al. identified seven polyporusterones from the fruiting bodies
of P. umbellatus and named them as A, B, C, D, E, F and G (Ohsawa et al., 1992).
Four new compounds were isolated from the sclerotia, 9α-hydroxy-1,2,3,4,5,10,19-
heptanorergosta-7,22-diene-6,9-lactone and ergosta-7,22-diene-3β,5α,6β-triol (Ohta
et al., 1996b) as well as 5α,8α-epidioxy-(24S)-24-methylcholest-6-en-3β-ol and
5α,8α-epidioxy-(24R)-24-methylcholesta-6,9(11),22-trien-3β-ol (Ohta et al., 1996a).
In addition, polyporusterones A and B previously recognized by Ohsawa et al.
(Ohsawa et al., 1992) were identified from the sclerotium by Ohta et al. (Ohta
et al., 1996a). Three alkaloids and two steroids were isolated from the sclerotia
of P. umbellatus; the structures of these were ascertained as 9-β-D-
ribofuranosyladenine (adenosine), 1-β-D-ribofuranosyluracil (uridine), 2,4-
pyrimidinedione (uracil), ergosta-4,6,8(14),22-tetraen-3-one and ergosta-5,7,22-
triene-3β-ol (ergosterol) on the basis of spectroscopic data and chemical correlations
(Lee et al., 2002). Two new polyporusterones named as polyprosteroneI and
polyprosterone II were isolated from sclerotia of P. umbellatus. Their structures
have been established based on spectral analysis (Zheng et al., 2004). Two other
polyperusterones named (20S,22R,24R)-16,22-epoxy-3β,14α,23β,25-tetrahydroxyergost-
7-en-6-one and (23R,24R,25R)-23,26-epoxy-3β,14α,21α,22α-tetra-hydroxyergost-
7-en-6-one were isolated from the sclerotia of P. umbellatus (Zhou et al., 2007).
Three polyporusterones were rediscovered in those experiments (Zhou et al.,
2007). Three new ecdysteroids (polyporoids A, B, C) with five known steroids,
which were previously identified by Ohsawa et al. (Ohsawa et al., 1992) and
Ohta et al. (Ohta et al., 1996b) were isolated from the ethyl acetate extract of
the sclerotium. All these ecdysteroids exhibit anti-inflammatory activity
against 12-O-tetradecanoylphorbol-13-acetate (TPA) induced inflammation in
mice. The inhibitory effects of ecdysteroids are higher than indomethacin, a
commercially available anti-inflammatory drug (Sun & Yasukawa, 2008). A new
pentacylictriterpene named 1-β-hydroxylfriedelin was recently isolated from the
P. umbellatus sclerotia and its structure has been elucidated (Zhao et al., 2009c).
Zhao et al. identified eight steroids in P. umbellatus using the HPLC coupled with
mass spectroscopy detection (Zhao et al., 2010c). Apart from (22E,24R)-ergosta-
6-en-3?,5?,6?-triol steroid, all other steroids were identified (Lu et al., 1985;
Ohsawa et al., 1992; Lee et al., 2002; Sun & Yasukawa, 2008). Ergone can be used
as a marker, in order to standardize production of P. umbellatus sclerotium. Since
ergone combines florescence properties, it can be easily admitted for quantitative
and qualitative analysis (Yuan et al., 2003). Quantitative analysis of ergone levels
in sclerotia have been carried out using a high performance liquid
chromatography-ultraviolet detector (HPLC-UV) (Yuan et al., 2003; Yuan et al.,
2004) and high performance liquid chromatography-fluorescence detector
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom: a review 11
(HPLC-FLD) and the results were verified using high performance liquid
chromatography-atmospheric pressure chemical ionization-mass spectrometry
(HPLC-APCI-MS/MS) (Zhao et al., 2009f; Zhao et al., 2009d). Ergone content
depends on a number of factors, such as genetic variation, fungus origin, drying
process and storage conditions (Zhao et al., 2009f). The HPLC-APCI-MS method
has been developed for qualitative analysis of known steroids in P. umbellatus,
and can therefore be used for the quality control of P. umbellatus. This is
necessary due to variations in the quality of P. umbellatus samples obtained from
different localities (Zhao, 2009; Zhao et al., 2010c). Zhao et al. subsequently
developed a more accurate and precise method to determine ergone
concentration from biological fluids using HPLC dual wavelength UV (Zhao
et al., 2010e). They developed a fast and sensitive HPLC–APCI-MS/MS method
for the determination of ergosta- 4,6,8(14),22-tetraen-3-one (ergone) in rat
plasma, the absolute recoveries of both ergone and ergosterol from the plasma
being more than 95% (Zhao et al., 2010b). The developed method has been
successfully applied to pharmacokinetic study of the drug in rats. Zhao et al.
introduced rapid resolution liquid chromatography with atmospheric pressure
chemical ionization tandem multi-stage mass spectrometry (RRLC-APCI-MSn)
and HPLC-FLD for identification and quantification of ergonefrom rat plasma,
urine and faeces (Zhao et al., 2010d). These methods are suitable for analysis in
preclinical and pharmacokinetic studies of ergone which is a major bioactive
component of P. umbellatus.
Zhao et al. developed RRLC-APCI-MSnand HPLC-APCI-MS/MS
method for the identification and quantification of ergosterol and its metabolites
in rat plasma, urine and faeces (Zhao et al., 2011a). Chen et al. carried out a
quantitative analysis of ergosterol recovered from blood plasma, urine and faeces
caused by orally administering ergosterol obtained from P. umbellatus (Chen et
al., 2013). The cloud-point extraction technique was used for the first time in
extracting ergosterol from blood plasma, urine and faeces, whereas HPLC-UV
was used for quantitative measurement. The results indicated that the ergosterol
level in faeces was higher than in plasma and urine (Chen et al., 2013). It was
shown that the above method is more suitable for pharmacokinetic analysis
carried out using ergosterol.
Introduction for medicinal properties
Polyporus umbellatus is commonly used in traditional Chinese medicine
(Huang & Liu, 2007; You et al., 1994; Zhao et al., 2009e). It was referred to in the
well-known medical book Shen Nung Pen Tsao Ching between A.D. 25-220 about
1,600 years prior to the earliest foreign record (1801) (Zhao & Zhang, 1992).
According to Li Shi-chen’s Compendium of Materia Medica, P. umbellatus
“opens up the texture and interspaces of the skin, and muscle, including the sweat
pore, cures gonorrheal swelling, beriberi, leucorrhea, gestational urination,
disturbances, foetus swelling and difficulty in urination” (Ying et al., 1987; Jong &
Birmingham, 1990).
Ying et al. recorded a number of Chinese traditional herbal formulas
including lignicolous mushrooms as sclerotia of P. umbellatus, which can be used
to treat conditions such as acute nephritis, systemic dropsy, thirst, difficulty in
urination, edema, urination disturbance, sunstroke, watery diarrhea, jaundice,
cirrhosis and ascites (Ying et al., 1987). It has a diuretic effect on pathogenic
dampness and is being used in traditional medicine combinations to treat oliguria,
12 A. R. Bandara et al.
edema, diarrhea, strangury with cloudy urine or leucorrhea (Wu, 2005; Liu & Liu,
2009). The sclerotium of P. umbellatus is also a Traditional Chinese Medicine
used for edema and promoting diuresis (Xiaoke & Shunxing, 2005; Xing et al.,
2012). Ecdysteroids, which exist in the sclerotia, act as a defensive mechanism and
exhibit various biological activities including in vitro cytotoxic, in vivo antitumor-
promoter, and antioxidant activities (Sun & Yasukawa, 2008; Ueno et al., 1980).
Presently the wild sclerotium of P. umbellatus is the main source for medicinal
uses (Xiaoke & Shunxing, 2005; Xing et al., 2013b).
Antitumor activity
Tumors, also known as neoplasms, are swellings or abscesses formed by
an abnormal proliferation of cells. Tumors can be benign, pre-malignant or
malignant (De Silva et al., 2012a). Ito et al. reported that water soluble glucan
from sclerotia of P. umbellatus demonstrated a strong antitumor activity against
subcutaneously implanted sarcoma 180, and also inhibited the growth of Shionogi
carcinoma 42 and pulmonary tumor 7423 in mice (Ito, 1973). Chemical analysis
confirmed this glucan-contains β-(1→3), β-(1→4), and β-(1→6) linked branches
and signified coexistence of β-(1→3) and β-(1→6) branches as indispensable for
the antitumor effect (Ito, 1973). Ito et al. investigated the influence of the sex of
experimental animals on the antitumor activity of polysaccharide from sclerotia
of P. umbellatus (Ito et al., 1975). It was observed that the growth rate of male
mice bearing Sarcoma 180, Ehrlich solid carcinoma, pulmonary tumor 7423 and
MF-sarcoma was higher than female mice of the same kind. In addition, the
regression rate of female mice treated with polysaccharides was high when
compared to the male mice. Both males and females which experienced a
regression of ascites tumor due to the administration of polysaccharides rejected
the re-implanted Ehrlich ascites carcinoma, Sarcoma 180, NF-sarcoma and
Shionogi carcinoma 42 (Ito et al., 1975).
Miyazaki et al. explained the structure of the antitumor glucans and
proposed the probable structural units (Miyazaki et al., 1978). Chemical analysis
of the antitumor glucans of sclerotia of P. umbellatus revealed that
polysaccharides bear the above mentioned linkages and it was further discovered
that β-(1→3)and β-(1→6) linkages were consistent in the backbone of the
structure of these glucans, while β-(1→4) and β-(1→6) linkages were found in the
branches connected with the backbone (Miyazaki et al., 1978). These glucans
cause complete regression of subcutaneously implanted sarcoma 180 tumor cells
in mice (Miyazaki et al., 1978). Miyazaki et al. disclosed that the basic common
unit of glucans of sclerotia from P. umbellatus is of primary importance and also
that the chemical structure of the glucans influenced antitumor activity (Miyazaki
et al., 1979). This activity was influenced by the type of sugar linkage, length of
the branch, branching frequency, molecular size and molecular conformation. The
probable structural units of the four antitumor glucans using P. umbellatus were
determined (Miyazaki et al., 1979). Ueno et al. isolated an alkali-soluble β-D-
glucan polysaccharide from sclerotia, similar to water soluble polysaccharides
composed of a backbone of β-(1→3)-linked D-glucopyranosyl residues, and
possessing of a single β-D-glucopyranosyl group joined through O-6 of every third
D-glucopyranosyl residue of the backbone (Ueno et al., 1980). In 1981, unknown
authors from Japan obtained a patent for an antitumor glucan, which was isolated
from the mycelium of P. umbellatus. This glucan was active against sarcoma 180
(Anonymous, 1981). Several alkali-soluble polysaccharides were isolated by Ueno
et al. who confirmed occurrence of (1→3)-β-D-glucan and found that the O-6
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom: a review 13
substituent in the (1→3)-β-D-glucan was of major importance in antitumor
activity against sarcoma 180. It was revealed that C-6 branched (1-3)-β-D-
glucopyranosyl-(1-3)-β-D-glucopyranosyl is the common unit of antitumor active
glucans and the branching frequency of the glucans is also significant for the
antitumor activity process (Miyazaki, 1983). It was shown that P. umbellatus
polysaccharides act against sarcoma 180 and Ehlich’s carcinoma tumor, using the
experiments carried out upon mice (Wang et al., 1983). It was shown that these
polysaccharides have the potential to suppress spontaneous metastasis in Lewis
lung sarcoma, and act against uterine cancer U14 (Wang et al., 1983).
Although Ito et al. signified that no activity was found in alkali soluble
glucans against tumor cells, rather than water-soluble glucans (Ito, 1973), Ueno
et al. showed that alkaline soluble polysaccharides are more effective than some
water-soluble polysaccharides. The authors confirmed that alkaline soluble
polysaccharides act against sarcoma 180 tumor cells more than water-soluble
polysaccharides (Ueno et al., 1982). Polysaccharides of P. umbellatus are known
for their adoptogenic antitumor effect on mice bearing hepatoma H22 (Wei et al.,
1983b; Wei et al., 1983a). The number of hepatoma H22cells in mice was reduced
after treatment and the plasma corticosterone and liver glycogen content as well
as enzyme activities were restored (Wei et al., 1983b; Wei et al., 1983a; Wu et al.,
1982). Polysaccharides with antitumor and immunomodulating activities have
been obtained from the liquid culture medium of P. umbellatus. A branched
D-glucotetraose was identified by Ogawa and Kaburagi as the repeating unit of
the extracellular polysaccharide of P. umbellatus (Jong & Birmingham, 1990).
Ito et al. gave a descriptive explanation of the mechanism of P. umbellatus
polysaccharides against Sarcoma 180 tumor cells in mice. It was shown that these
polysaccharides do not exhibit a direct cytocidal action against the tumors, while
the activation of the C3, stimulation of the reticuloendothelial system and the
inhibition of hepatic drug metabolizing enzymes cause a direct cytocidal action
(Ito, 1986). Polyporus umbellatus combined with mitomycin C enhanced the life
span of mice with an intrahepatic implantation of sarcoma 180 tumor cells by
inhibiting the synthetic rates of DNA, RNA and protein in tumor cells (You et al.,
1994). Cachexia, a common condition in many human cancer patients,
particularly in gastrointestinal or lung cancer patients, is characterized by loss of
weight, muscle atrophy, fatigue and weakness (Muscaritoli et al., 2006). Cachexia
results in eventual death of these patients. Toxohormone-L, a protein that inhibits
food and water intake promoting anorexia was found in the patients with
cachexia. P. umbellatus polysaccharides reduced cachexia caused by
toxohormone-L protein in rats (Wu et al., 1997). Xu et al. investigated cytotoxic
activity of peripheral blood monocytes which is activated by polysaccharides of
P. umbellatus and Interleukin 2 against tumor cells in culture. Cytotoxicity of
killer cells co-stimulated by the polysaccharides and Interleukin 2 which are
effective against both natural killer resistant and natural killer sensitive tumor
cells (Xu et al., 1998). Polyporus umbellatus polysaccharide enhances cytotoxic
activity mediated by natural killer cells against target cells of YAC-1 cells and
P-815 cells of mice (Nie et al., 2000).
Chen et al. manufactured a tablet composed of components from
P. umbellatus, Poria cocos and skin pulp of two Bufo species. They reported
potential antitumor, immunostimulant and analgesic properties and the ability to
remove toxic substances. This mixture has been used to produce pharmaceuticals
and foods, which helps the human body against radiation and chemical (Chen
et al., 2007). The compound extract constituted P. umbellatus and two other
14 A. R. Bandara et al.
traditional Chinese medicines Agrimonia pilosa and Gambogia (dry resin secreted
by Garcinia hanburyi) and inhibited human gastric carcinoma MGC-803 tumor
cell growth in vitro and in vivo in a dose dependent manner. In an experiment
carried out upon a sample of mice, it was shown that this compound induces
programmed cell death in the above tumor cells and may be a promising novel
anti-tumor drug in human gastric carcinoma (Zhao et al., 2009a). According to an
analysis done using different Chinese Traditional Antitumor Medicines, i.e.,
Ligustrazine Hydrochloride, Astragalus Mongholicus Bge, Matrine N-Oxide and
Artesunate, P. umbellatus polysaccharides exhibit a down regulating effect upon
immunosuppressors of colorectal tumor cells in vitro (Li et al., 2011).
Other than polysaccharides, ergone extracted from the sclerotium of
P. umbellatus shows an outstanding antitumor effect against human hepatocellular
carcinoma HepG2 cells (Zhao et al., 2011b). Cell proliferation is inhibited due
to the effect of these tumor cells, upon the G2/M phase of the cell cycle and
the induction of apoptosis generated from the caspase activation The above
mentioned GUMP-1-1 and GUMP-1-2 polysaccharides (Bi et al., 2013)
significantly brought down tumor volumes in hepatoma H22 transplanted mice.
These two polysaccharides maximum tumor inhibition rate and maximum life
prolonged was recorded at a dose of 200mg/kg. Bi et al. also demonstrated that
GUMP-1-1 and GUMP-1-2 P. umbellatus polysaccharides indicate a significant
antitumor activity (Bi et al., 2013).
Anticancer activity
The early stage of cancer is referred to as neoplasm, and exhibits
uncontrolled cell proliferation resulting in an abnormal mass of cells (De Silva
et al., 2012a). Later these cells spread to surrounding tissues and even to distant
sites. Cancers always show a malignant growth of cells (De Silva et al., 2012a).
Polyporus umbellatus is used as a medicine, especially in anticancer drugs (Wei
et al., 1983b; Zhao & Zhang, 1992). Polyporus umbellatus sclerotium have figured
prominently in Chinese pharmacopeia, especially in the treatment of lung cancer
(Ying et al., 1987; Stamets, 2000). A polysaccharide extract (Khz) obtained fused
mycelia of P. umbellatus and Ganoderma lucidum, inhibits the growth of A549
lung cancer cells (Kim et al., 2012). Yang et al. pointed out experimentally and
clinically that P. umbellatus inhibits the induction of bladder cancer (Yang, 1991).
A clinical study evaluated the prophylactic effect of P. umbellatus on bladder
cancer. It was shown that the stimulating immune responses of P. umbellatus and
Bacillus Calmette Guerin on bladder recurrence was better than mitocycin C
(Yang et al., 1999). All seven polyperostones isolated from the fruiting body of
P. umbellatus showed cytotoxic action on leukemia 1210 cell lines and inhibited
cell proliferation (Ohsawa et al., 1992). Histopathological studies showed that
lymphocytes infiltrated and surrounded the cancer cells, and there was fibrosis
in both normal and cancerous cells. These results indicate the potential use of
P. umbellatus as an anticancer agent (You et al., 1994). Polyperostone A and B
show greater cytotoxicity in higher concentrations (Sekiya et al., 2005). Methanol
extracts of sclerotium of P. umbellatus exhibited a cytotoxic effect against human
gastric cancer cells and ergone inhibited the growth of cancer cell lines in colon,
cervix, liver and stomach. The cytotoxic effects were stronger against cancerous
cells in liver and colon, than the cervix and stomach cancer cells (Lee et al., 2005).
Ergone, (22E,24R)-ergosta-7,22-dien-3β-ol, 5α,8α-epidioxy-(22E,24R)-
ergosta-6,22-dien-3β-ol, ergosta-6,22-dien-3β,5α,6β-triol, and polyporusterone B
which were isolated from P. umbellatus show anticancer activity against HepG2,
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom: a review 15
Hep-2, and Hela cancer cells, while ergone exhibits selective cytotoxic activity
against cancer cells (Zhao et al., 2010a). Aqueous extracts of sclerotia of
P. umbellatus are highly effective in inhibiting bladder carcinogenesis in rats,
which is also associated with up regulation of glutathione S-transferase πand
NAD(P)H quinoneoxidoreductase 1 in the bladder (Zhang et al., 2011).
Immune system enhancement
Extracts of P. umbellatus, in a drug produced by “Institute of Chinese
drugs, Academy of Chinese Traditional Medicine”, enhance immunity (Ying et al.,
1987). It was reported that the polysaccharides of P. umbellatus show a significant
protective effect against acute toxicity in mice livers and the treatment
reestablished the activities of liver glucose-6-phospate and acid phosphatase (Lin
& Wu, 1988). Polysaccharides of P. umbellatus produce significant hepatoprotective
activity against hepatotoxicity caused by CCl4and D-galactosamine in mice (Lin &
Wu, 1988). Zhang et al. showed that in normal mice as well as in mice which had
liver lesions upon using CCl4; the number of macrophages and the amount of
H2O2released in peritoneal cavities were increased by polysaccharides from
P. umbellatus (Zhang et al., 1991). Polysaccharides from P. umbellatus can boost
the cellular immunity of both normal mice and those with liver lesions (Zhang
et al., 1991). It was shown that P. umbellatus polysaccharides enhance the
lymphocyte function of immunosuppressed mice, and upregulate the number of
CD4+ T cells and IgG level as reported. The authors also concluded that
P. umbellatus polysaccharide acts as an immune response upregulator (Nie et al.,
2000).
Yang et al. investigated the immunosuppressive effects of culture
supernatant of sarcoma 180 cells in the presence or absence of P. umbellatus
polysaccharide. The study was carried out using mice, where it was shown that
P. umbellatus polysaccharides have the potential to offset the immunosuppressive
effects taking place due to culture supernatant of sarcoma 180 cells, as well as
downregulate the immunosuppressive substances which are synthesized and/or
secreted by the culture supernatant of sarcoma 180 cells (Yang et al., 2004). It was
shown that the polysaccharides extracted separately from P. umbellatus mycelium
and the sclerotium using aqueous extracts advanced the weight of immunological
organs when administered orally to mice. It was shown that these two
polysaccharides are similar and that no significant difference was observed in their
ability to advance the weights of immunological organs (Tian et al., 2005). Li et al.
administrated orally to mice the polysaccharides extracted separately from
P. umbellatus mycelium and sclerotium using an aqueous extract and followed this
up with the celiac mononuclear macrophage test, erythrocyte rosette formation
test, metatarsal swelling thickness test, lymphocyte transformation test, and EAC
rosette test. It was established that the test group differed significantly (P< 0.05)
from the control group, with P. umbellatus polysaccharides increasing the
immunity of white mice (Li et al., 2007).
Pan et al. investigated the possibility of irradiation prevention and
immunity regulation of P. umbellatus polysaccharides in vitro and in vivo.
Polyporus umbellatus polysaccharides amplified cell proliferation rate and
CD43+cell count of umbilical cord blood hematopoietic stem cells culture in vitro.
Mice with transplanted umbilical cord blood hematopoietic stem cells and treated
with P. umbellatus polysaccharides had the lowest death rate and shortest
recorded recovery time. These polysaccharides amplify the hematopoietic stem
cells, and these cells promote the immune and hematopoietic reconstruction of
16 A. R. Bandara et al.
transplanted mice (Pan et al., 2008). Li et al. showed P. umbellatus polysaccharides
induced phenotypic and functional changes in murine bone derived dendritic
cells via toll-like receptor 4 (TLR-4). Polyporus umbellatus polysaccharides
significantly stimulate the proliferation of mouse splenocytes and upregulated the
expression of CD86 and CD11c in a dose dependent manner. The polysaccharide
induces dendritic cell maturation and differentiation and then activates natural
killer and Th1 cells to enhance immune responses. Polyporus umbellatus
polysaccharides could also activate CD4+CD45RA+ T cells (Li et al., 2010; Li &
Xu, 2011a). A novel study also showed aqueously extracted polysaccharides from
fermented mycelium of P. umbellatus increased the killing potency of natural
killer cells of mouse spleen and promoted proliferation of mouse B and T cells
(Sun & Zhou, 2014). Li and Xu studied the molecular mechanism of its
immunostimulatory potency and immune responses of macrophages, using
polysaccharides prepared from aqueous extract of P. umbellatus. The aqueous
extract upregulated the activity of macrophages, while stimulating splenocyte
proliferation and production of cytokines, as well as cytotoxic and inflammatory
molecules. From experiments carried out using mice, it was concluded that the
polysaccharides of P. umbellatus cause the increment of immune stimulating
potency via TLR-4 activation of the signaling pathway (Li & Xu, 2011b).
Water-soluble polysaccharides extracted from the fruiting body of
P. umbellatus have the potential to activate B cells, macrophages and dendritic
cells. Depletion of branches of the polysaccharides causes a substantial reduction
in the ability not only to activate B cells in vitro, but also to elicit specific IgM
production in vivo. Virtually all healthy human subjects possess high-titer
circulating antibodies that work against the ZPS backbone, suggesting that ZPS
epitope is shared by environmental antigens capable of eliciting adaptive humoral
responses in the population (Dai et al., 2012). β-Glucans are major polysaccharide
constituents of P. umbellatus (Miyazaki & Oikawa, 1973; Jong & Birmingham,
1990) and considered to be valuable biological response modifiers for their ability
to enhance the activity of immune cells, aid in wound healing and prevent
infections (Dai et al., 2012). Polysaccharides extracted from P. umbellatus possess
immunomodulatory activities (Peng et al., 2012). Aqueously extracted
polysaccharides, GUMP-1-1 and GUMP-1-2 could remarkably increase the spleen
weight and splenocyte proliferation of hepatoma H22 tumor bearing mice, as a
consequence improve the immune response (Bi et al., 2013). These results indicate
that the P. umbellatus immune activities are most probably due to its
polysaccharides.
Diuretic effect
Sclerotia of P. umbellatus have been used from a long time in Traditional
Chinese Medicines for urological disorders (Zjawiony, 2004; Sekiya et al., 2005;
Zhao et al., 2009f). They are prominently used as herbal remedy with or without
the combinations of other medications, in order to treat patients suffering from
chronic kidney diseases (Zhao et al., 2012b). In particular, an aqueous extract of
dried sclerotia is traditionally used for diuresis (promoting urination) (Ying et al.,
1987; Jong & Birmingham, 1990; Yuan et al., 2004; Wu, 2005; Zhao et al., 2010e;
Xing et al., 2012).
The sclerotia of P. umbellatus is considered as an urination promoting
component in traditional Chinese formulas such as Gorei-san (), Chorei-to
(), Irei-to ()Bunsyou-to (), and Inchingorei-san
(), (Yuan et al., 2004; Zhao et al., 2009f) which promote the diuretic
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom: a review 17
process and govern water metabolism. This signifies a relative diuretic effect
causing reinforcement of water pathway functions capable of draining dampness,
as is remarked in the ancient work Shennong’s Herbal Classics ().
Furthermore, the sclerotium boosts the urination process and prevents dampness
which in turn prevents further results in preventing edema, scanty urine, vaginal
discharge, cloudy painful urinary dysfunction, jaundice and diarrhea (Wang et al.,
1964; Ying et al., 1987; Yuan et al., 2004; Xing et al., 2012; Zhao et al., 2012b).
Clinical studies have confirmed that P. umbellatus is an effective diuretic medicine
for the treatment of pyelonephritis, nephritis and urologic calculi without side
effects (Jyothi, 2013).
Wang et al. demonstrated the diuretic effect of P. umbellatus by
administrating decoction of sclerotium to un-anesthetized dogs, which increased
urine output and excretion of sodium, potassium, and chloride ions (Wang et al.,
1964). Through experiments done with mice, it was concluded that the anti-
aldosteronic effect of ergosta-4,6,8(14),22-tetraen-3-one (ergone) contained in
P. umbellatus sclerotia promotes urination (Yuan et al., 2004). Ergone isolated
from many mushrooms has been shown to possess antialdosteronic diuretic
properties (Lindequist et al., 2005) and proven to prevent progression of renal
injury and subsequent renal fibrosis (Zhao et al., 2012a).
In oral dozes of P. umbellatus administered in Chinese Traditional
Medicine treatments, (Wu, 2005) ergone is absorbed via the oral route (Yuan
et al., 2004). Although it was previously shown that ergone does not significantly
affect urinary sodium and potassium excretion in normal rats (Yuan et al., 2004),
Zhao et al. demonstrated that ergone increased potassium, sodium and chloride
excretion and the volume of urine excreted in normal rats (Zhao et al., 2009e).
In addition to ergone, the ergosterol and D-mannitol components included in
P. umbellatus facilitate the diuretic process. Ergoneis the strongest diuretic drugs
among these components (Zhao et al., 2009e).
A patent has been issued for ergone as a diuretic drug (Zhao et al.,
2009b). Ergone contained in P. umbellatus, normalizes nitrogenous products and
regulation of ions in unbalanced blood and urine of mice. Early administration of
ergone prevents progression of renal injury and subsequent renal fibrosis in
aristolochic acid nephropathy (Zhao et al., 2011c). Although it is time dependent,
with a sharp difference in metabolic profile, ergone present in P. umbellatus aids
recovery from chronic renal failure in rats (Zhao et al., 2012b). Kawashima et al.
introduced a Choreito Japanese Traditional Medicine, which comprises sclerotia
of P. umbellatus, as a successful treatment for renal disorders (Kawashima et al.,
2012).
Antioxidant and free radical scavenging activity
Free radical-induced oxidation is affective in the pathological processes
which cause prolonged and degenerative diseases, such as cardiovascular disease,
cancer, diabetes, neurodegenerative disease and ageing (Gan et al., 2010). These
free radicals damage cells causing DNA mutations, protein inactivation, lipid
peroxidation, and cell apoptosis (Gan et al., 2010). Mushrooms play a significant
role in the search for efficacious, non-toxic substances, with free radical
scavenging activity because the protein content of the polysaccharide extracts has
a direct effect on free radical scavenging activity (Liu et al., 1997). Due to the high
protein content in the polysaccharide extract of fruiting body of P. umbellatus, it
exhibits a strong superoxide and moderate hydroxyl radical scavenging activity
when compared to Coriolus versicolor, Ganoderma lucidum, Lentinula edodes,
18 A. R. Bandara et al.
Schizophyllum commune, Tremella fuciformis, Trichloma lobeyense and
Volvariella volvacea (Liu et al., 1997). These studies signal that the anti-oxidant
activity of P. umbellatus polysaccharides have the potential to treat oxidative
stress related diseases.
Anti-oxidative activity plays an important role in atherogenesis,
inflammation and ageing (Sekiya et al., 2005). It was shown that P. umbellatus
exhibits anti-oxidant and free radical scavenging activity in human red blood
cells (RBC) in vitro and in vivo in mice (Sekiya et al., 2005). 2,2-azo-
bis(2-amidinopropane)dihydrochloride, free radical initiator induces hemolysis of
RBC, while aqueous extracts of P. umbellatus inhibit this activity. Further they
demonstrated that the two triterpenes, polyporusterones A and B present in a
aqueous extract of P. umbellatus, exhibit inhibitory activities against free radical
induced hemolysis of red blood cells in vitro. The antioxidative effect was dose-
dependent and P. umbellatus strengthens the antioxidative effect of plasma in
vivo. Results of in vivo tests indicated that P. umbellatus inhibited the free radical
generation dose dependently. The free radical scavenging activities of a group of
rats, treated P. umbellatus were significantly higher than the control group (Sekiya
et al., 2005).
Gan et al. evaluated the anti-oxidant activity and total phenolic contents
of Chinese medicinal plants including P. umbellatus, which are used in treating
rheumatic diseases. These evaluations were deduced using ferric-reducing
antioxidant power and Trolox equivalent antioxidant capacity assays, and the
values obtained for P. umbellatus were lower than other plants. They showed that,
the phenolic compounds generate major increases in anti-oxidant capacity, while
the polysaccharides of P. umbellatus showed anti-oxidant activity, although the
phenolic content of P. umbellatus is lower (Gan et al., 2010).
One of the antioxidant mechanisms of P. umbellatus was demonstrated
using experiments on mice livers. Polysaccharides of P. umbellatus are able to
suppress hepatic lipid peroxidation by increasing hepatic malondialdehyde: a
major reactive aldehyde that is formed in the degradation of polyunsaturated
lipids catalyzed by reactive oxygen species (Nielsen et al., 1997). These
polysaccharides also increased hepatic levels of glutathione which is an
endogenous non-enzymatic antioxidant and major antioxidant such as superoxide
dismutase, glutathione peroxidase, catalase in carbon tetrachloride treated mice
together with the up regulation of their mRNA expression (Peng et al., 2012).
Free radical activity of the recently isolated GUMP-1-1 and GUMP-1-2
polysaccharides was investigated using a P. umbellatus aqueous extract (Bi et al.,
2013). During investigation, GUMP-1-2 exhibited a strong scavenging activity
upon hydroxyl and superoxide free radicals, while in GUMP-1-1it was weak. The
scavenging effect of GUMP-1-2 was dose dependent and exhibited significant
increment on superoxide free radicals at a concentration of 0.8mg/ml. The high
ironic acid content and average molecular weight of GUMP-1-2 are causes of this
antioxidant activity (Bi et al., 2013).
Hair growth
Polyporus umbellatus significantly increases regrowth of hair of mice
(Inaoka et al., 1994). 3,4-dihydroxybenzaldehyde extracted from sclerotium of
P. umbellatus has a high potential to stimulate the regrowth of hair (Inaoka et al.,
1994). Ishida et al. identified hair regrowth promoting compounds as
polyporusterones A, B which were previously isolated by Ohsawa et al. (Ohsawa
et al., 1992) and a new compound, acetosyringone (Ishida et al., 1999b). Among
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom: a review 19
these, polyporusterone A is said to be more effective in mammals and the crystal
structure has been analyzed further (Ishida et al., 1999a).
Anti-viral activity
Combined administration of polysaccharides from P. umbellatus and
Salvia miltiorrhizae increases normalization rate of alanine transaminase and
negative conversion rate of HBeAg in patients with chronic hepatitis B (Xiong,
1993). Polysaccharides of P. umbellatus have the potential to cure chronic
hepatitis B virus. These polysaccharides induce an effect upon the clearance of
serum hepatitis B antigen and hepatitis B virus DNA and thereby present a
possible cure for chronic hepatitis B (Liu et al., 2001). Due to immune modulatory
effects, polysaccharides of P. umbellatus have been widely used to treat hepatitis
B or C together with antiviral drugs in the form of injections or tablets in China
(Peng et al., 2012). Hao et al. introduced a traditional Chinese medicine, which
includes polysaccharides of P. umbellatus, plant and fungi ingredients and bears
a higher curative rate and rapid action against AIDS. According to the authors,
it is a non-toxic drug with no side effects (Hao et al., 2011). Zhan patented a
traditional Chinese medicine with extracts of P. umbellatus and related herbs,
which inhibits the reproduction of the HIV and improves the CD4 immunocyte
level (Zhan, 2012). Zhang et al. discovered a traditional Chinese medicine which
improves the CD4+T lymphocyte level against the AIDS virus and exhibits good
anti-inflammatory and abirritation effects (Zhang et al., 2012c).
Anti-bacterial activity
Polyporus umbellatus is used in China as an antibacterial drug (College,
1982). Polyporus umbellatus exhibits strong inhibitory activity in vitro against
urogenital Chlamydia trachomatis (Li et al., 2000), the most common bacterial
sexually transmitted disease (Black, 1997). Extract of fermentation broth of
P. umbellatus shows antibiosis against Bacillus subtilis, Candida tropicalis,
Escherichia coli, Fusarium graminearum, Saccharomyces cerevisae, and
Staphylococcus aureus (Sun & Zhou, 2014; Wang et al., 2009). This bacteriostasis
substance is similar to non-water-soluble type II antibiotics and sensitive to acid
alkali and unstable to heat. This poorly stable antibiotic-like substance and ester-
peptide antibiotic shows similar absorption pattern in UV spectrum (Wang et al.,
2009).
Anti-protozoal activity
Polyporus umbellatus showed inhibitory activity against protozoan
parasite Plasmodium falciparum, one of the main causative agents of malaria in
humans (Lovy et al., 2000).
Cultivation
Cultivation for sclerotium production
Due to the perceived medicinal value of P. umbellatus, the commercial
need for sclerotia has greatly increased in recent years. The result is that the wild
source of P. umbellatus will soon be exhausted (Guo et al., 2002). The decrease in
wild sclerotial production and the increase in demand have stimulated interest
in the search for substitutes for the natural source of sclerotium (Liu & Guo,
2009). However, to face the high demands of the global market, it is necessary
20 A. R. Bandara et al.
to cultivate strains of P. umbellatus under artificial or semi artificial growth
conditions (Huang & Liu, 2007; Zhou et al., 2007).
Sclerotia of P. umbellatus were successfully cultured in China co-
inoculated with A. mellea (plantation, 1978). This technique is not useful for large
scale production due to slow growth and the difficulty of obtaining sufficient seed
sclerotia from nature for artificial cultivation (Guo et al., 2002). Guo and Li
reported that the hyphae isolated from basidiospores of P. umbellatus successfully
formed white or brown sclerotia in solid and liquid medium (Guo & Li, 1982).
Wang et al. also obtained sclerotia in liquid medium (Wang et al., 1982). However,
the methods were limited to laboratory conditions and unable to meet the
requirements for mass production (Guo et al., 2002). Guo and Xu developed a
technique for cultivating sclerotia of P. umbellatus (Guo & Xu, 1993). Guo et al.
produced sclerotium in an artificial media using a dual culture method (Guo et al.,
2002). They demonstrated that P. umbellatus could not form sclerotia without
A. mellea. Xing and Guo artificially developed the sclerotium of P. umbellatus in
a wheat bran medium and showed that the artificially developed and wild sclerotia
are morphologically very similar (Xiaoke & Shunxing, 2005). The studies
concluded that P. umbellatus favours aerobic conditions and therefore the burying
depth of inoculum plays a significant role in its cultivation (Choi et al., 2003).
Sclerotia cultivated using root inoculation develop more quickly than those
cultivated when buried. Root inoculation has been found more appropriate for
the development of the sclerotia of P. umbellatus due to many beneficial factors
such as the simplicity of the inoculation process, reduced cultivation period and
facility of harvest (Choi et al., 2003). Yang cultivated P. umbellatus in a sawdust-
based medium. It is a method which has both a brief productive cycle and a high
survival rate, and could be tried at the industrial level (Yang, 2003).
Liu patented a method for cultivating P. umbellatus using basswood
dibbling and an implanting method or rot plant embedding method. These
methods enable the sclerotium to develop and immediately form fruiting bodies
without the support of any other companion fungus. The strong resistance,
impurity repulsing ability, lack of seasonal limitation and cheapness are significant
benefits of this method (Liu, 2004). Guo et al. patented three growth media where
sclerotial formation equivalent to that of wild sclerotia obtained from the mycelia
of P. umbellatus are produced; they proposed these as industrially useful methods
with high sclerotia formation ability (Guo et al., 2007).
It was found that P. umbellatus sclerotium could be proliferated with high
efficiency in a short period of time, through symbiotic culturing with A. mellea
(Kikuchi, 2007). The method of cultivating P. umbellatus with A. mellea in the
natural environment under applicable environmental conditions improves the
quality and yield of artificially cultured P. umbellatus (He et al., 2007). A low cost
method which can rapidly produce the sclerotia of P. umbellatus - using corn grits
and wood chips and/or blocks of media inside polythene bags - was developed (Jin
et al., 2010). Zhang introduced a simple, low cost method of growing P. umbellatus
in humic acid media with prepared natural organic substances (Zhang, 2011). Lee
et al. introduced a method for the cultivation of P. umbellatus using agricultural
and industrial by-products but without pesticides and heavy metals and avoiding
the use of soil (Lee et al., 2011). A method of inter-planting Glastrodia elata with
P. umbellatus, after culturing A. mellea with Glastrodia elata, was reported (Sun
et al., 2011). Xue introduced an artificial cultivation method by imitating the
growing mechanism of the wild sclerotia. This method generated a higher
production rate in a short period of time and produced better-purified sclerotia
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom: a review 21
(Xue, 2012). A large-scale cultivation method aimed at producing higher yields,
using a tank type pit inside forests was described by Zhang (Zhang, 2012b).
The carbon source (the type of medium used in producing P. umbellatus
sclerotia artificially) is significant in the induction of sclerotium formation (Cheng
et al., 2006). It was shown that for the formation of sclerotia an appropriate
medium is malt extract agar modified with GPC (Glucose, Peptone, Corn steep
liquor), and 18-25° optimum temperature (Cheng et al., 2006). It was confirmed
that fructose and peptone were the best carbon and nitrogen sources for
sclerotium formation (Liu & Guo, 2009). The carbon source affects the formation
of sclerotia, while the nitrogen source influences morphological transformation.
Vitamins and minerals are not essentially needed for the sclerotial formation (Liu
& Guo, 2009). Xing et al. highlighted that the carbon source and an initial high pH
are essential factors for sclerotial formation at low temperature in sawdust media
(Xing et al., 2013b). They concluded that mycelia subjected to environmental
stress by exposure to low temperatures and enhanced reactive oxygen species can
induce higher sclerotial formation and polysaccharide content than in nutritional
agar medium (Xing et al., 2013b). Presently cultivation of P. umbellatus is being
carried out in China, through artificial infection of Armillaria (Kikuchi, 2007;
Kikuchi & Yamaji, 2010; Xing et al., 2012; Zhou et al., 2012; Xing et al., 2013b).
Cultivation for mycelium production
Suitable carbon and nitrogen sources for mycelial growth and
extracellular polysaccharides production are glucose and yeast extract (Gu et al.,
2001). Of six carbohydrates, fructose, glucose, sucrose and starch significantly
promoted mycelial growth and starch was most effective for the production of
mycelium (Lee et al., 2007). A submerged culture media was optimized by Lee
et al. for production of ergone using mycelium of P. umbellatus, wherein the
mycelium and ergone production were significantly increased by co-culturing
P. umbellatus with A. mellea (Lee et al., 2007). Cui et al. discovered a method to
increase the yield of mycelium of P. umbellatus in a short time, by pre-fermenting
the growing media with A. mellea. This is a low cost method which produces high
amounts of polysaccharides and minimizes heavy metal contents (Cui et al., 2007).
Guo et al. patented a low cost medium which contains wheat bran and glucose that
boosts higher mycelium yield and polysaccharide content (Guo et al., 2008).
Huang and Liu investigated the optimum conditions required for the growth of
P. umbellatus mycelium, and for production of exopolysaccharides. They observed
that glucose and yeast extracts are the best carbon and nitrogen sources, and pH5
and 6 are optimum (Huang & Liu, 2008). Xing et al. investigated the
environmental factors, using optimum mycelium growth of P. umbellatus at pH8
in dark conditions and temperature of 25°C and a maximum polysaccharide
content was produced at pH8 –10 at 5°C (Xing et al., 2012).
Although yeast and peptone were found to be the best nitrogen sources
for P. umbellatus, the costs are prohibitive. Therefore their use for fermentation
on an industrial scale is not viable (Chen et al., 2010a). Chen et al. used a
submerged culture fermentation method, using whey as a cheap alternative
nitrogen source which facilitated higher mycelium growth and high
exopolysaccharide production. The maximum biomass and exopolysaccharides
production obtained was from 3% glucose and 50% whey broth (Chen et al.,
2010a). Table 1 lists some of the methods used for cultivation and production of
polysaccharides from P. umbellatus.
22 A. R. Bandara et al.
Table 1. Methods used for cultivation of sclerotia and production of polysaccharide derivatives from P. umbellatus.Abrreviations: asl = above sea
level, eps =exopolysaccharide, mat=mean annual temperature
Type
of the method Materials Host used for
experiment Yield Time taken Mode References
With support of
Armillaria sp.
1Both mycelia (P. umbellatus and A. mellea)
plugs were cultured in
Sawdust wheat bran medium in flask
grit medium in flower pots Quercus variabilis
Sclerotia
30 days
90-120 days
Laboratory (Guo et al., 2002;
Xiaoke & Shunxing,
2005)
2Sclerotia attached lateral root of host and
A. tabescens, or A. melleapre-inoculated
wood logs
Castanea crenata
Quercus mongolica Sclerotia weight
increased
7-40 times
10 months Outdoor (Choi et al., 2003)
3Sclerotia with A. mellea or Armillaria sp.
pre-inoculated wood logs None Sclerotia weight
not increased
significantly
12 months Outdoor (Choi et al., 2003)
4A. mellea inoculated sticks with P. umbellatus
sclerotia slices as seeds + tree leaves in a dug
pit (70 cm ×30 cm) in forest. Cellar covered
by soil
Broad leaved wood
stick (unknown sp.)
Diameter 8-12 cm,
length 60 cm
Sclerotia yield
unknown Unknown Scrub forest
1000-1800 m, asl,
mat.11-12°C,
soil pH 5-6.5
(He et al., 2007)
51
st stage: Corn flour, sucrose or glucose, beef
extract, water and agar
2nd stage: 1st stage culture + A. mellea corn
grits, wood chip and/or wood block
3rd stage: Sub cultured 2nd stage in
polypropylene bagwith2nd stage culture
medium
Broad leaved wood
(unknown sp.) Sclerotia yield
unknown 30-45 days 20-25°C
(Jin et al., 2010)
6Culturing of A mellea, in G. elata seeds, sowing
P. umbellatus and those seeds layer by layer G. elata Sclerotia yield
unknown Unknown Outdoor (Sun et al., 2011)
7Sand, wood chips with A. mellea pre-
inoculated wood and P. umbellatus in boxes +
liquid fertilizer
Unknown wood sp. Yield unknown Unknown Indoor (under
controlled
temperature and
humidity)
(Fan, 2014)
8Materials (bagasse or corncob media + wheat
bran, corn powder, lime, and/or sugar, water)
pre fermented with A. mellea and inoculated
P. umbellatus in plastic bags
Unknown eps yield
unknown 5-7 months Indoor18-32°C (Cui et al., 2007)
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom: a review 23
9Starch, fructose, peptone, formic acid,
KH2PO4, MgSO4, FeSO4, yeast extract,
media 25 °C, pH 4.5(liquid medium)
None Dry mycelium
yield 3.5 g/l,
Ergone 86.9 µg/g
of dry mycelium
15 days Laboratory
25°C (Lee et al., 2007)
Without
support of
Armillaria sp.
1Sclerotia initially formed on PDA and after
growing on sawdust bran medium None Sclerotia max
weight 30 g 25 days on
PDA, about
70-90 days on
sawdust
Indoor (Yang, 2003)
2Cultured on PDA and then cultivated on
sawdust, bran, rice sweets, and chaff medium None Fruiting body (Liu, 2004)
3Cultured on PDA
Next in a liquid media
Then in three solid media as follows.
1. glycerol, peptone, corn steep liquor, agar,
water
2. mannitol, peptone, corn steep liquor, agar,
and water
3. sawdust (or wheat stalk, corn stalk,
or corn cob), soybean cake powder, glycerol,
corn steep liquor, and water
None Sclerotia yield
unknown 30-50 days
12-20 days
25-50 days
25-50 days
30-120 days
(Guo et al., 2007)
4Fructose 50.0 g/l, peptone 4.0 g/l, K2HPO4
1 g/l, KH2PO40.46 g/l, MgSO40.5 g/l, vitamin
B10.05 mg/l, agar 10 g/l, deionized water
5.40 g of sclerotial
weight/100 g
substrate
30-40 days Laboratory (Liu & Guo, 2009)
5Sawdust, cottonseed hull, brewers grain etc.
medium cultured in a box, a paper bag or a
bottle on the ground
Yield unknown Unknown Indoor (Lee et al., 2011)
6Glucose 3.5%, peptone 3.0%, yeast extract
0.2%, KH2PO40.3%, MgSO40.15%, and
vitamin B10.001%) + P. umbellatus broth
concentrate 5-10% (pH 5.5) (liquid medium)
eps production
310 mg/ml 36 hours Laboratory
25°C (Zhou et al., 2001)
Table 1. Methods used for cultivation of sclerotia and production of polysaccharide derivatives from P. umbellatus.Abrreviations: asl = above sea
level, eps =exopolysaccharide, mat=mean annual temperature (continued)
Type
of the method Materials Host used for
experiment Yield Time taken Mode References
24 A. R. Bandara et al.
7Wheat bran, glucose, KH2PO4, MgSO4
(liquid medium) Myceliumeps
yield unknown 18-30 days Laboratory (Guo et al., 2008)
8Main media (glucose 2.5%, peptone 0.5%,
yeast extract 0.5%, KH2PO40.1%, MgSO4:
7H2O 0.1%, and vitamin B10.005%) +
P. umbellatus broth concentrate7 (pH 5.5)
(liquid medium)
Mycelia
production 12.7 g/l 11 days Laboratory
(rotary shaker
at 25°C, 100 rpm)
(Huang & Liu, 2007)
9glucose 3%, skim milk 0.2%, KH2PO40.1%,
MgSO4: 7H2O 0.1%, and vitamin B10.005%
(pH 5)(liquid medium)
eps production
0.571 g/l 14 days Laboratory
(rotary shaker
100 rpm, at 25°)
(Huang & Liu, 2008)
10 glucose 3%, whey broth 50%, KH2PO40.1%,
MgSO4: 7H2O 0.1%and vitamin B10.005%
(liquid medium)
eps production
0.632 g/l 14 days Laboratory
(rotary shaker at
25°C, 100 rpm)
(Chen et al., 2010a)
Unknown
(with or without
Armillariasp.)
1P. umbellatus (sclerotia as seeds) +humic acid
media 5-7 parts + primary soil 3-4 parts by
weight in dredging cellar (1.3-1.6 m ×1-1.3 m
×1-1.3m)
Sclerotia yield
unknown Dredging cellar
built in slope land
(15-35°C)
(Zhang, 2011)
2Tank type pit in forest
Size of pit 30 cm ×80 cm ×3 m Sclerotia can be
harvested more
than one time
Dwarf shrub forest
land 1,000-1,500 m
asl
(Zhang, 2012b)
Table 1. Methods used for cultivation of sclerotia and production of polysaccharide derivatives from P. umbellatus.Abrreviations: asl = above sea
level, eps =exopolysaccharide, mat=mean annual temperature (continued)
Type
of the method Materials Host used for
experiment Yield Time taken Mode References
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom: a review 25
Products
Due to the above mentioned medicinal effects, the edible mushroom
P. umbellatus is used as a non-toxic and low cost bioactive ingredient for
manufactured pharmaceutical products, and food supplements, having no side
effects as well as in cosmetics and beverages
Medicinal products and dietary supplements
Tian et al. manufactured an injection composed of antitumor
polysaccharides of P. umbellatus. Less than 3000 patients were used for clinical
testing of this product and no significant side effects were recorded (Tian et al.,
1980). Kim and Lee produced a medicine incorporating P. umbellatus which
inhibits immune-modulation and tumor growth. This medicine promotes the
T-cells activity, increases production of interferon-γand interleukin-6 and inhibits
angiogenesis and tumor growth (Kim & Lee, 2006). An antitumor drug with
P. umbellatus and other herbs which has synergistic effects and can be made into
oral formulations and injections was produced (Wei, 2007). Li and Wang
manufactured a probiotic with P. umbellatus polysaccharides, which improved
immunity in tumor bearing mice and nutritional conditions of cancer patients (Li
& Wang, 2009). Tien-Hsien liquid (THL), which is used as an anti-cancer dietary
supplement is a herbal mixture of P. umbellatus extracts and 13 other Chinese
herbs. The antitumor and cytotoxic properties of P. umbellatus cause induction of
apoptosis of human cancer cells (Sun et al., 2005). MycoPhyto Complex (Plate 1)
is an anticancer dietary supplement, which includes a blend of mycelia of
P. umbellatus and five other mushroom species. It has potential therapeutic value
in the treatment of invasive human breast cancer (Jiang & Sliva, 2010). Using
P. umbellatus, a pill with anti-tumor and anti-aging effects was introduced by
Chen et al. (Chen et al., 2012). Zhang manufactured a tablet with 30-80%
P. umbellatus polysaccharides which minimizes adverse effects and maintains fast
absorption. This tablet is able to adjust immunity function and can be used as
adjuvant medicine in bladder cancer mice (Zhang, 2012a).
Cho et al. developed a red ginseng extraction product including
P. umbellatus, which effectively improves sexual enhancement with powerful
ejaculation (Cho et al., 2004). Kuok et al. manufactured a herbal product which
includes P. umbellatus as an ingredient preventing and treatments of prostate
disorders including prostatitis, benign prostate hyperplasia, prostatic carcinoma,
tumor, elevated blood levels of prostate specific antigen and irritative voiding
symptoms such as nocturia and excessive frequency and urgency of urination
(Kuok & Ly, 2004). A medicinal composition which contained P. umbellatus is
used for treating gynecological inflammation such as pelvic inflammation, chronic
pelvic inflammatory disease, ovarian cystitis, colpitis, cervicitis and adnexitis.
Apart from relieving inflammation it can be used to stop bleeding, and release
smooth muscle spasms in the uterus (Wang & Hou, 2007).
A food supplement which includes P. umbellatus and some other
components was produced by Takeda, and was used as a treatment for high
cholesterol, diabetes, hypertension, and liver problems (Takeda, 2005). The
product of Cho et al. has been effective also in preventing arteriosclerosis,
hypertension and diabetes (Cho et al., 2004). Similarly, Chai et al. manufactured
a health care product, which can improve hypoglycemic, hypolipidemic and
immunity (Chai et al., 2012).
A herbal extract including P. umbellatus which can repress the level
of acetaldehyde in blood was produced; this product represses the level of
acetaldehyde in alcoholic digestion and heals hangover (Kim et al., 2010).
26 A. R. Bandara et al.
A herbal drink, with P. umbellatus was introduced by Hong which alleviated
alcohol hangover and recovered the liver function (Hong, 2010). Li introduced a
beverage, which can relieve alcoholism and nourish the liver (Li, 2012).
Wang and Wang obtained a patent for Chinese medicine compositions
which include P. umbellatus, and these productions show therapeutic effect upon
digestive system disorders and nausea, vomiting, dizziness, and hypotension
(Wang & Wang, 2010).
Qu produced a pill, which includes P. umbellatus polysaccharides,
for treatments of Hepatitis B (Qu, 2005). A sulfate compound, containing
P. umbellatus and with an anti-hepatitis B virus activity, was synthesized (Liu
et al., 2006). A Chinese medicine composition, which includes P. umbellatus has
been used for treatments of hepatitis and fatty liver (Zhang, 2007). A medicinal
composition with P. umbellatus can be effective in promoting blood circulation
and diuresis, relieving pain, killing Escherichia coli and Candida albicans (Wang
& Hou, 2007). Miao et al. produced an immune protection agent against fowl
infectious bursal disease virus, using P. umbellatus (Miao et al., 2010).
Market research and development is taking place on P. umbellatus
capsules, injections, lyophilized powder agent and many preparations. Zhang et al.
registered a Chinese patent for manufacturing powder and polysaccharides using
the P. umbellatus mycelium as pharmacological products, i.e., granules, capsules
and oral liquids (Zhang et al., 2005). The capsule disintegration is slow and thereby
the biological use is said to be low; Injection is in frozen-dry powder form and
production costs are high and therefore the medicine is not cost effective. Tablets
with high speed disintegration and dispersion are more convenient and have high
biological use (Zhang et al., 2012a). Patent certifications have been issued to many
Chinese and Korean medicinal compositions which contain P. umbellatus as a
major component. These medicinal compositions are used to treat many human
diseases. Accordingly, P. umbellatus is used in Chinese medicinal compositions for
treatments of condyloma (Qin, 2013), Obesity (Fang et al., 2012; Ran, 2013; Wang,
2013a; Wang, 2013b), abdominal distention caused in the early stages of Traumatic
injury, relieving constipation and swelling of limbs (Xia et al., 2013), common cold
in young children (Zhang et al., 2013a), the treatments and prevention of facial
paralysis (Liu, 2013), throat pain and oral ulcers (Sun, 2013b), diabetes mellitus
(Ding & Dai, 2013; Lee et al., 2013; Sun, 2013a; Sun, 2014), glomerulonephritis
(Guo, 2012), chronic nephritis (Li, 2013a), pyelonephritis (Yan et al., 2013), the
treatments and prevention of liver related diseases such as fatty liver (Sun, 2013c;
Zhou, 2013), chronic hepatitis and cirrhosis (Fu et al., 2013; Hu & Lu, 2013; Xu,
2013; Zhao et al., 2013), the treatments of chronic renal failure (Wang, 2013c),
nephropathy (Guo, 2013), diarrhea (Ma, 2013b; Li et al., 2013), leg ulcers (Qiu &
Teng, 2013), infantile bronchial asthma causing due to cold fluid- retention and
fever (He & Xie, 2013), inflammation and immune dysfunction caused by multi
resistant bacteria infections (Kim, 2012; Zhang et al., 2013b), urinary stone (Li,
2013c), damp-heat type gallstone (Ma, 2013a), chronic enteritis (Zou, 2013), acute
mastitis causing due to alcoholism (Yuan, 2013), acute mastitis due to milk stasis
(Xing, 2013), chronic renal insufficiency (Xin, 2013), hepatitis B (Li, 2013b),
nodular prurigo (Zeng, 2013), cystitis (He, 2013; Yuan et al., 2012), macular
hemorrhage of high myopia (Guan, 2013), hydroperitoneum (Shao & Wu, 2013),
eczema (Zhan & Zhan, 2013), senile retinopathy and macular degeneration (Hu et
al., 2013), dysmenorrhea and uterine bleeding (Wang, 2014), benign prostatic
hyperplasia and urinary disorders such as urinary urgency, difficulty urinating,
incontinence, urinary retention, hematuria (Zeng, 2014) Dietary supplements which
contain P. umbellatus and their beneficial effects are shown in Table 2 and Plate 1.
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom: a review 27
Table 2. Dietary supplements with P. umbellatus alone or in mixtures. The co-authors of the present paper have not confirmed these claims
Name Category Tablet/Capsule Doze Ingredients Function Certified by Web page
Complexe
Hepato Bio Dietary
supplement 1000 mg
Capsules 1 to 2
capsules
per day
P. umbellatus
mycelium 20% Enhance liver function Certified according
to EU Reg. 834/07 http://www.nature-et-forme.com/
complexe-hepato-bio/p466#info_tabs_1
Mushroom 6
Immune
Support
Complex
Capsules 1 to 2
capsules
1-3 times
daily
Cordyceps sinensis,
Ganoderma lucidum,
Lentinula edodes,
Poria cocos,
P. umbellatus,
Trametes versicolor,
Relief of inflammatory
conditions, supporting
immune function,
supporting white blood
cell production and
providing antioxidant
protection
http://www.bioceuticals.com.au/
product/preview/Mushroom-6/print
Mycophyto®
Complex Dietary
supplement Powder/
Capsules 4g-8g or
6 capsules/
1-3 times
daily
Agaricus subrufescens,
Coriolus, Ganoderma,
Cordyceps sinensis,
P. umbellatus, Maitake
Strength immune system http://www.choosecra.com/store/
supplements/myco-phyto.html
Polyporus-
MRL Dietary
supplement 500 mg
Tablet P. umbellatus
mycelium and
primordia
Supports the immune
system 100%Organic in the
USA by Quality
Assurance International
and EU Council
Regulation (EEC)
No. 2092/91
http://www.mycologyresearch.com/
products.asp?product=
Polyporus
P. umbellatus,
180 capsules 300 mg
Capsules Up to
3capsules
per day
P. umbellatus fruitbody antitumor effects,
diuretic effects,
antioxidant and free-
radical scavenging
activity, immune system
enhancement, hair
growth, antiviral effects
Good manufacturing
practices quality
assured
http://www.shopssl.de/epages/
es105220.sf/en_GB/?ObjectPath=/
Shops/es105220_Prime-Visions/
Products/PU180
P. umbellatus
Extract 500 mg
Capsules P. umbellatus
sclerotium Enhancesthebody’s immune
response to an antigen http://www.activehealth.co.uk/Polyporus-
Umbellatus-Extract__p-70.aspx
Polyporus
Fungal Body Dietary
supplement 1000 mg
Capsules P. umbellatus mycelium
Poria cocos mycelium Unknown http://dsld.nlm.nih.gov/dsld/
Ingredient.jsp?db=adsld%2C&item=
POLYPORUS+FUNGAL+BODY
P. umbellatus –
HdT Food
supplement 450 mg
Capsules 1-2 capsule
per day Suitable for diabetics and
people who suffer from
celiac disease
http://www.hifasdaterra.com/index.php/
extracto-ecologico-polyporus-hdt/
Zhu Ling
(P. umbellatus)
Hot-water extract
425 mg
Capsules P. umbellatus
sclerotium Diuretic, anticancer
activity http://www.mushroomnutrition.com/
polyporus/polyporus-mn-60-cab.html
28 A. R. Bandara et al.
Cosmetics
Polyporus umbellatus is used for production of cosmetics. Tsuji et al.
produced topical and bath preparations composed of P. umbellatus that prevent
skin aging (Tsuji et al., 1995), A product including P. umbellatus extracts inhibits
testosterone 5α-reductase activity and antimicrobial activity against
propionibacterium acnes, which can be used for acne prevention and treatment
(Rang et al., 2009), A herbal product of P. umbellatus with antioxidant, cell-
activating, collagen synthesis-promoting effects for skin ageing prevention was
produced (Yoon et al., 2012), β(1→6) branched β(1→3) glucan as an active
ingredient of P. umbellatus can deter skin aging, impart skin whitening effect and
cure skin damage effectively (Du et al., 2014; Hyde et al., 2010). The β-glucan was
also found promising as an active ingredient in anti-wrinkle activity, wound
healing, antioxidant activity and moisturizing effect (Du et al., 2014; Hyde et al.,
2010). Therefore P. umbellatus has a cosmetics producing potential.
Food and beverages
Due to the promising and enduring effects of P. umbellatus as a
medicinal mushroom, it is used in the manufacture of various foods and
beverages. Wine produced with extracts of P. umbellatus as an ingredient, is a
healthy drink, which includes amino acids and vitamins (Zhou et al., 2008). A wine
(Baek et al., 2012a) and a rice extracts (Baek et al., 2012b) which contained,
mycelial extract of P. umbellatus bare anti-diabetic and anti-obesity effects. Yoon
manufactured a sauce containing P. umbellatus (Yoon, 2012). Han produced an
Fig. 1. Examples of Polyporus umbellatus – containing products. 1. ComplexeHepato Bio;
2. Mushroom 6 Immune Support Complex; 3. Myco-Phyto Complex; 4. Polyporus-MRL;
5. Polyporus umbellatus-HdT; 6. Polyporus umbellatus Extract; 7. Zhu Ling (Polyporus
umbellatus) Hot water extract.
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom: a review 29
instant tea powder consisting of edible mushrooms including P. umbellatus,
claiming the potential to prevent respiratory tract infections (Han, 2013).
A product of tea granules containing P. umbellatus is capable of nourishing blood
and prevent anemia (Liu, 2014). Hot water preparations containing P. umbellatus
are health improving beverages (Lee et al., 2012).
Other polyporus umbellatus-containing products
Polyporus umbellatus is used in the production of fertilizers: Wu
produced a fertilizer for the cultivation of Daucus carota using P. umbellatus
(Wu, 2010) while Xu developed a fertilizer for the cultivation of Arctium lappa
using P. umbellatus as a component (Xu, 2010).
Many suppliers who have been selling extract of P. umbellatus advertise
online. Most of them are Chinese suppliers (Alibaba.com). Experiments proved
that the polysaccharides content of these extracts vary between 10% and 40%.
Fruiting body and sclerotium of P. umbellatus were used as source and extraction
was done using solvents such as hot water and ethanol resulting in a final greyish
brown powdery product. Supplying capability varies from 200kg to 200 tons per
month. Under proper storage conditions these powders have a long shelf life, up
to 2 to 3 years (Alibaba.com).
CONCLUSION AND PERSPECTIVES
Mushrooms have long been valued by the mankind as a medicinal
resource (De Silva et al., 2012a; De Silva et al., 2012b; De Silva et al., 2013;
Poucheret et al., 2006; Wasser, 2002). Mushrooms or their extracts are used
globally in the form of dietary supplements (Jiang & Sliva, 2010). Indeed, fungi
form a major and largely untapped source of powerful new pharmaceutical
products (Lo & Wasser, 2011; Poucheret et al., 2006; Wasser, 2002) Polyporus
umbellatus contains biologically active substances in cultured mycelium, cultured
broth, fruit body and sclerotium. P. umbellatus has the potential to promote
diuretic action, and as a medicinal treatment for many chronic and serious
diseases. For example, P. umbellatus has the potential to treat cancers, which are
the second largest cause of death in people (Daba & Ezeronye, 2003), HIV,
Chlamydia trachomatis – the most common sexually transmitted diseases in the
United States (Black, 1997) and Diabetes mellitus – causing 2.2 % of deaths in
the world (De Silva et al., 2012b; Lo & Wasser, 2011). In addition, P. umbellatus
has shown anti-obesity (Baek et al., 2012b) and anti-skin ageing (Tsuji et al., 1995)
properties without any side effects. Therefore with potential medicinal value
P. umbellatus has great value in the global market (Huang & Liu, 2007).
Polyporus umbellatus can be used to produce exopolysaccharides and
ergone. Exopolysaccharides produced from mushrooms have been shown to have
special medical effects in clinical trials. In addition, polysaccharides can also be
used in industrial applications such as emulsifying and foam stabilizing agents,
food coatings and thickening agents (Chen et al., 2010b).
Polyporus umbellatus is one of the most valuable medicinal mushrooms
and widely used in east Asian countries such as China, Japan, Korea and Taiwan
(Huang & Liu, 2007; Yin et al., 2012). Nowadays the demand on P. umbellatus has
increased drastically due to its promising effects (Zhou et al., 2007). Taking Korea
30 A. R. Bandara et al.
as an example, the demand for P. umbellatus has increased year on year since they
began to use it as an herbal medicine and as a result they now import it from
China (Choi et al., 2002). Wild sources of P. umbellatus are seriously depleted due
to a lack of effective protection (Xing et al., 2013b; Huang & Liu, 2007; Xiaoke &
Shunxing, 2005) and over-exploitation due to demand on the global market (Yin
et al., 2012). It is therefore considered as an endangered medicinal fungus in China
(Zhang et al., 2012b).
In practice, it would take a long time to cultivate P. umbellatus in both
solid and submerged cultivations (Chen et al., 2010b). The fungus has a long lag
phase and mycelial growth of P. umbellatus is much slower than that of other
mushroom species (Huang & Liu, 2007). Artificial cultivation of P. umbellatus is
time consuming and labour intensive (Huang & Liu, 2007). Cultivation of
P. umbellatus via infection with A. mellea has been practiced over the past
30 years; this technique is restricted by a low proliferation rate, unstable yield and
lack of natural sclerotia to serve as seeds (Xu et al., 1998).
It is an unsolved problem that the sclerotium is not produced directly
from the hyphae, which effectively impedes the production scale and the
production efficiency of P. umbellatus (Xiaoke & Shunxing, 2005). It is necessary
to develop efficient artificial cultivating methods for developing the sclerotia and
fruiting bodies within shorter time periods to meet demand in the global market.
Asexual propagation is signified as the main pathway followed in order to
produce cultivated products of P. umbellatus (Zhang et al., 2010a).
On the other hand P. umbellatus can be used sustainably by reducing
overexploitation and preventing the depletion of natural habitats. Polyporus
umbellatus grows successfully in forest ecosystems, while forest management
practices such as tree cutting (specially host trees) interrupts the growth of the
former (Kunca, 2011). Polyporus umbellatus is able to produce new sclerotia
under appropriate conditions. Due to this it is possible to enhance the natural
production by conserving natural habitat. It is also possible to make people aware
that during the harvesting of sclerotia retaining some as seeds will allow the
microhabitat to be reconstructed. Depletion of natural habitats undoubtedly
checks the presence of this mushroom (Yin et al., 2012).
Polyporus umbellatus is an edible medicinal mushroom of great interest
for healthy people or patients mainly used by food, pharmaceutical and cosmetic
industries. Many products are indeed developed from P. umbellatus mycelium,
sclerotium and exopolysaccharides such as natural health foods and traditional
medicine as well as food supplements used to prevent, support or cure several
diseases.
Acknowledgements. We would like to thank Humidtropics, a CGIAR Research
Program that aims to develop new opportunities for improved livelihoods in a sustainable
environment, for partially funding this work. Kevin D. Hyde thanks the Chinese Academy
of Sciences, project number 2013T2S0030, for the award of Visiting Professorship for
Senior International Scientists at Kunming Institute of Botany. INRA (Project AGASIA
of the regional program Bio-Asie), the project Value added products from Basidiomycetes:
Putting Thailand’s biodiversity to use (BRN049/2553), the National Research Council of
Thailand (NRCT), projects – Taxonomy, Phylogeny and cultivation of Lentinus species in
northern Thailand (NRCT/55201020007), Mae Fah Luang University project – Taxonomy,
Phylogeny and cultivation of Lentinus species in northern Thailand (MFU/54 1 01 02 00 48),
and Thailand Research Fund grant – Taxonomy, Phylogeny and biochemistry of Thai
Basidiomycetes (BRG 5580009) are also thanked for supporting this study.
Polyporus umbellatus, an Edible-Medicinal Cultivated Mushroom: a review 31
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