Fomitopsis betulina (formerly Piptoporus betulinus): the Iceman’s polypore fungus with modern biotechnological potential

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World J Microbiol Biotechnol (2017) 33:83
DOI 10.1007/s11274-017-2247-0
REVIEW
Fomitopsis betulina (formerly Piptoporus betulinus): theIceman’s
polypore fungus withmodern biotechnological potential
MałgorzataPleszczyńska1· MartaK.Lemieszek2· MarekSiwulski4·
AdrianWiater1· WojciechRzeski2,3· JanuszSzczodrak1
Received: 31 January 2017 / Accepted: 15 March 2017
© The Author(s) 2017. This article is an open access publication
a promising source for the development of new products for
healthcare and other biotechnological uses.
Keywords Biological activity· Cultivation· Fomitopsis
betulina· Phytochemistry· Piptoporus betulinus
Introduction
In 1991, a mummified body was discovered in the Val Sen-
ales glacier in Italy. The man (named Ӧtzi the Iceman), who
lived 5300 years ago, carried two fragments of a fruiting
body of Fomitopsis betulina (formerly Piptoporus betuli-
nus). Some scientists believe that Ӧtzi might have used the
fungus for medical purposes (Capasso 1998) and, although
the idea arouses some controversy (Pöder 2005), the long
tradition of the use of F. betulina in folk medicine is a fact
(Reshetnikov etal. 2001; Wasser 2010). Infusion from F.
betulina fruiting bodies was popular, especially in Russia,
Baltic countries, Hungary, Romania for its nutritional and
calming properties. Fungal tea was used against various
cancer types, as an immunoenhancing, anti-parasitic agent,
and a remedy for gastrointestinal disorders (Grienke etal.
2014; Lucas 1960; Peintner and Pöder 2000; Semerdžieva
and Veselský 1986; Shamtsyan etal. 2004). Antiseptic and
anti-bleeding dressings made from fresh F. betulina fruit-
ing body were applied to wounds and the powder obtained
from dried ones was used as a painkiller (Grienke et al.
2014; Papp etal. 2015; Rutalek 2002).
In the present paper, we have shown the current knowl-
edge of the fungus F. betulina, including its lifestyle, chem-
ical composition, and potential in biotechnology.
Abstract Higher Basidiomycota have been used in natural
medicine throughout the world for centuries. One of such
fungi is Fomitopsis betulina (formerly Piptoporus betuli-
nus), which causes brown rot of birch wood. Annual white
to brownish fruiting bodies of the species can be found on
trees in the northern hemisphere but F. betulina can also be
cultured as a mycelium and fruiting body. The fungus has a
long tradition of being applied in folk medicine as an anti-
microbial, anticancer, and anti-inflammatory agent. Proba-
bly due to the curative properties, pieces of its fruiting body
were carried by Ötzi the Iceman. Modern research confirms
the health-promoting benefits of F. betulina. Pharmacologi-
cal studies have provided evidence supporting the antibac-
terial, anti-parasitic, antiviral, anti-inflammatory, antican-
cer, neuroprotective, and immunomodulating activities of
F. betulina preparations. Biologically active compounds
such as triterpenoids have been isolated. The mushroom is
also a reservoir of valuable enzymes and other substances
such as cell wall (1→3)-α-d-glucan which can be used for
induction of microbial enzymes degrading cariogenic den-
tal biofilm. In conclusion, F. betulina can be considered as
* Małgorzata Pleszczyńska
m.pleszczynska@poczta.umcs.lublin.pl
1 Department ofIndustrial Microbiology, Maria Curie-
Skłodowska University, Akademicka 19, 20-033Lublin,
Poland
2 Department ofMedical Biology, Institute ofRural Health,
Jaczewskiego 2, 20-095Lublin, Poland
3 Department ofVirology andImmunology, Maria Curie-
Skłodowska University, Akademicka 19, 20-033Lublin,
Poland
4 Department ofVegetable Crops, Poznań University ofLife
Sciences, Dąbrowskiego 159, 60-594Poznań, Poland

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Taxonomy andcharacteristics
Piptoporus betulinus (Bull.) P. Karst. (known as birch poly-
pore, birch bracket, or razor strop) is a common Basidi-
omycota brown rot macrofungus growing on decaying
birch wood. Homobasidiomycetes were divided into eight
clades. The family Polyporaceae with the genus Piptopo-
rus was classified to the polyporoid clade, and then the
antrodia clade—the Fomitopsis-Daedalea-Piptoporus
group comprising brown rot fungi was identified within
this clade (Hibbett and Donoghue 2001; Hibbett and Thorn
2001). Further studies of the phylogenetic relationships
among members of the antrodia clade revealed polyphyly
of the Fomitopsis genus and suggested that P. betulinus
was phylogenetically closer to Fomitopsis than to Piptopo-
rus (Kim etal. 2005; Ortiz-Santana etal. 2013). Recently,
P. betulinus (Bull.) P. Karst. has been transferred to Fomi-
topsis (Han etal. 2016) and, according to Index Fungorum
(2016), is classified in the genus Fomitopsis, family Fomi-
topsidaceae, order Polyporales, class Agaricomycetes, divi-
sion Basidiomycota, kingdom Fungi, with the current name
Fomitopsis betulina (Bull.) B.K. Cui, M.L. Han and Y.C.
Dai, comb.nov. (MycoBank no.: MB 812646).
Fomitopsis betulina is characterized by annual, sessile to
effused-reflexed, tough to woody hard basidiocarps, white
to tan or pinkish-colored pore surface with mostly small
and regular pores. Fruiting bodies grow singly or in small
groups, are covered with a laccate, glabrous crust, never
zonate, young cream to white, later ochraceous-brown
to greyish brown (Fig.1a). The mycelium of F. betulina
developing on agar media is white, relatively homogene-
ous, downy-felt, with regular colony edges (Fig.1b). The
hyphae develop radially. The hyphal system is mostly dim-
itic. The clamped generative hyphae, 1.5–3.5µm in diam-
eter, are branched and hyaline whereas the skeletal hyphae
with the diameter of 3– 4 µm, are less branched and have
thicker walls. No primordia or fruiting bodies of this spe-
cies were found invitro (Petre and Tanase 2013). Basidi-
ospores are smooth, hyaline, thin-walled, and cylindri-
cal (Han and Cui 2015; Han etal. 2016; Kim etal. 2005;
Schwarze 1993).
The birch polypore grows mainly as a saprophyte on
dead trees and occasionally as a parasite of living trees. It
occurs in northern temperate forests and parks in Europe,
North America, and Asia. The host range of the fungus is
restricted exclusively to birch species, e.g. Betula pendula
Roth., B. pubescens Ehrh., B. papyrifera Marsh., and B.
obscura Kotula (Schwarze 1993; Žižka etal. 2010).
Wood decay
Wood rotting fungi are traditionally divided into white and
brown rot species based on the structure and composition
of residual wood. Brown rot fungi extensively degrade the
carbohydrate fraction of lignocellulose but, in contrast to
white rot fungi, leave lignin, although in a modified form.
In these fungi, chemical depolymerization of cellulose,
which precedes and supports its enzymatic degradation,
is very important. They lack ligninolytic peroxidases and
usually some other enzymes such as processive cellobio-
hydrolases used for degradation of crystalline cellulose,
but contain H2O2-generating oxidases and Fe3+- and qui-
none-reducing enzymes used for non-enzymatic depolym-
erization of polysaccharides (Arantes and Goodell 2014;
Baldrian and Valášková 2008; Hori etal. 2013). Modern
phylogenetic evidence suggest, however, that there is no
sharp distinction between the two groups of fungi (Hori
etal. 2013; Riley etal. 2014).
Fomitopsis betulina is one of the most common brown rot
species but its wood-decaying mechanism has been tested
only fragmentarily (Meng etal. 2012) and is still poorly
Fig. 1 Fomitopsis betulina. a Basidiocarp of the wild fungus. b Mycelium on an agar plate. c Mature fruiting body cultured on birch sawdust in
artificial conditions. (photographed by M. Siwulski)

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understood. As other fungi of this type, it degrades wood to
yield brown, cubical cracks easily broken down. Many fac-
tors, including microflora or compounds present in wood,
contribute to this complex process (Przybył and Żłobińska-
Podejma 2000; Song etal. 2016; Zarzyński 2009). Shang
etal. (2013) showed that wood samples decayed by F. betu-
lina lost 57% of dry weight (dw) and 74% of holocellulose
after 30 days, whereas the fungus growing on wheat straw
causes 65% loss of dw within 98 days of culture (Valášková
and Baldrian 2006a). A set of enzymes of F. betulina
involved in the degradation of lignocellulose was character-
ized in detail by Valášková and Baldrian (2006a, b). The
fungus growing on straw produced enzymes with wide sub-
strate specificities: (1→4)-β-endoglucanase, β-glucosidase,
(1→4)-β-endoxylanase, (1→4)-β-endomannanase,
(1→4)-β-xylosidase, and (1→4)-β-mannosidase. The
activities of ligninolytic enzymes and cellobiose dehydro-
genase for oxidoreductive cleavage of cellulose were not
detected. Similar results were obtained in liquid cultures by
Vĕtrovský etal. (2013). When F. betulina grew in nature,
β-glucosidase and β-mannosidase activity was associ-
ated with the fruiting bodies while endopolysaccharidases
were detected in colonized wood (Valášková and Baldrian
2006a).
Cultivation
Carpophores of F. betulina from natural habitats or myce-
lium and culture liquid from submerged cultures were used
as raw material to obtain extracts and bioactive substances
with medicinal properties (Table1) (Lomberh etal. 2002).
Studies concerning the mycelium growth rate in the pres-
ence of various substances (metals, dyes) were conducted
mainly on agar media or in liquid cultures (Baldrian and
Gabriel 2002; Dresch etal. 2015; Hartikainen etal. 2016).
The yield of F. betulina mycelium was established in liquid
cultures with addition of some agricultural wastes in the
studies of Krupodorova and Barshteyn (2015). The enzy-
matic activity of F. betulina was studied in laboratory con-
ditions on agar media (Krupodorova etal. 2014), in liquid
cultures (Vĕtrovský etal. 2013), on wheat straw (Valášková
and Baldrian 2006a, b), and on Betula sp. wood samples
(Reh etal. 1986; Shang etal. 2013).
There are limited data on small- or large-scale cultiva-
tion of this species in which carpophores could be obtained
in controlled conditions. The first such report referring to
outdoor log cultivation of F. betulina on Betula davurica
Pallas originated from Korea (Ka etal. 2008). Logs with
a diameter of 8–18 cm and length of 107–135 cm were
inoculated and then cultured in natural conditions. The
yield obtained was in the range from 212 to 1298g fresh
weight (1–2 mushrooms per log). Development of fruiting
bodies took an average of 18 months. The ratio of log yield
was estimated at 2.8–6.1%. The only report on indoor
production of F. betulina fruiting bodies was given by
Pleszczyńska etal. (2016). In the study, four strains of F.
betulina isolated from natural habitats were applied. Their
mycelia were inoculated into birch sawdust supplemented
with organic additives. Mature fruiting bodies weigh-
ing from 50 to 120g were obtained from only one strain,
after 3–4 months of the cultivation in artificial conditions
(Fig.1c). The biological efficiency ranged from 12 to 16%.
It was shown that extracts isolated from cultivated and
naturally grown F. betulina fruiting bodies had comparable
biological activity (Table1).
Biotechnological uses
Phytochemistry andpharmacological activity
Comprehensive analyses of the chemical composition of
the F. betulina fruiting body carried out under different
conditions (Grishin etal. 2016; Hybelbauerová etal. 2008;
Reis etal. 2011) revealed the presence of 17 fatty acids,
in it 22% saturated and 78% unsaturated (mainly oleic and
linoleic acid); sugars (d-arabinitol, d-mannitol and α,α tre-
halose); biomolecules with antioxidant properties (tocophe-
rols—0.578mg/100g dw, mainly β and γ; ascorbic acid—
87.5mg/100g dw; β-carotene and lycopene). Among other
identified compounds were betulinic acid, betulin, lupeol,
fomefficinic acid, ergosterol peroxide, and 9,11-dehydroer-
gosterol peroxide (Alresly etal. 2016; Jasicka-Misiak etal.
2010). Total content of phenolics was determined on 14 or
35mg GAE/g dw whereas phenolic acids were not detected
(Reis etal. 2011; Sułkowska-Ziaja etal. 2012). Product of
hydrodistillation of F. betulina fruiting bodies contained
numerous volatile mono- and sesquiterpenes. Several com-
pounds found, (+)-α-barbatene, (−)-β-barbatene, daucene
and isobazzanene, have not been previously reported from
other mushrooms. Alcohols, 3-octanol and 1-octen-3-ol,
were the main flavour constituents of the fungus (Rapior
etal. 1996; Rösecke etal. 2000).
Although some authors considered young specimens
of F. betulina edible (Wasson 1969), the fungus value
is not the result of nutritional but therapeutic proper-
ties. The overview of the available literature concern-
ing medical potential of birch polypore was presented
in Table1. Referring to the folk uses of the birch poly-
pore, most of the presented research was based on crude
extracts, which often have greater bioactivity than iso-
lated constituents at an equivalent dose. This phenom-
enon is explained by mostly synergistic interactions
between compounds present in mixtures. Furthermore,
extracts often contain substances that inhibit multi-drug

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Table 1 Biological properties of extracts and compounds isolated from Fomitopsis betulina
Biological activity Mechanism of biological activity Model [method of study] ExtractaActive compoundaReferences
Bactericidal Inhibition of bacterial growth Bacillus subtilis, Mycobacterium
smegmatis, Pseudomonas aerugi-
nosa, Serratia marcescens, Staphy-
lococcus aureus [zone of inhibition,
agar well diffusion assay]
Extracts Suay etal. (2000)
Brucella sp.[zone of inhibition, agar
well diffusion assay]
Benzene extracts Polyporenic acid (suggested) Utzig and Fertig (1957)
Bacillus sp., Rhodococcus equi, S.
aureus [zone of inhibition, disk-
diffusion method]
Chloroform extracts Karaman etal. (2009)
B. subtilis, Escherichia coli [zone of
inhibition, agar well diffusion assay]
Dichloromethane extracts Keller etal. (2002)
Bacillus sp., R. equi, S. aureus, E.coli
[zone of inhibition, agar well diffu-
sion assay]
Methanol extracts Karaman etal. (2009), Keller etal.
(2002)
B. subtilis, Sarcina lutea [zone of inhi-
bition, agar well diffusion assay]
Ethanol extracts Polyporenic acid A (suggested) Kandefer-Szerszeń etal. (1981)
B. subtilis, S. lutea, Brucella sp. [zone
of inhibition, agar well diffusion
assay]
Ether extracts Polyporenic acid (suggested) Kandefer-Szerszeń and Kawecki (1974),
Utzig and Fertig (1957)
B. subtilis, Enterococcus faecalis,
E.coli, S. aureus [zone of inhibition,
agar well diffusion assay, NCCLS-
method]
Piptamine isolated from submerged
culture of F. betulina Schlegel etal. (2000)
B. subtilis, E.coli, S. aureus [zone of
inhibition assay]
Mycelium, culture liquid Krupodorova etal. (2016)
B. subtilis, S. aureus [zone of inhibi-
tion assay]
3β-acetoxy-16α hydroxyl-24-oxo-5α-
lanosta-8-ene-21-oic acid
Alresly etal. (2016)
E. faecalis [zone of inhibition assay] Alkali extract Vunduk etal. (2015)
Fungicidal Inhibition of fungal growth Saccharomyces cerevisiae, Aspergillus
fumigatus, [zone of inhibition, agar
well diffusion assay]
Extracts Suay etal. (2000)
Candida albicans, Kluyveromyces
marxianus, Rhodotorula rubra,
Sporobolomyces salmonicolor,
Penicillium notatum [zone of
inhibition, agar well diffusion assay,
NCCLS-method]
Piptamine isolated from submerged
culture of F. betulina Schlegel etal. (2000)
Larvicidal Induction of larva death Aedes aegypti [bioassay] Dichloromethane extract Keller etal. (2002)
Antiviral Protection of CEF cells from vaccinia
virus Host/target cells: primary culture of
chick embryo fibroblast (CEF)
Challenge virus: vaccinia virus
Ethanol extracts Kandefer-Szerszeń etal. (1981)
Induction of sub stance with properties
similar to interferon (hot-stable,
stable at pH 2, nondialyzing, insensi-
tive to RNA-se, slightly sensitive to
trypsin)
[Plaque formation assays] Water extracts Kandefer-Szerszeń and Kawecki (1979)
Ether extracts polyporenic acid (suggested) Kandefer-Szerszeń and Kawecki (1974)
nucleic acids (RNA and DNA) Kandefer-Szerszeń etal. (1979)

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Table 1 (continued)
Biological activity Mechanism of biological activity Model [method of study] ExtractaActive compoundaReferences
Protection of HAT cells from vaccinia
virus by induction of interferon
Host/target cells: human fibroblast
culture (HAT)
challenge virus: vaccinia virus
[plaque formation assays]
RNA Kawecki etal. (1978)
Mice protection from lethal infection
with TBE
Host/target: Swiss mice
Challenge virus: tick borne encephali-
tis (TBE) virus
Ethanol extracts Kandefer-Szerszeń etal. (1981)
Water extracts induced substance with
properties similar to interferon (sta-
ble at pH 2, nondialyzing, sensitive
to trypsin)
[Neutralization test] Water extracts Kandefer-Szerszeń and Kawecki (1979)
Ether extracts Polyporenic acid Kandefer-Szerszeń and Kawecki (1974)
Nucleic acids (RNA and DNA) (sug-
gested)
Kandefer-Szerszeń etal. (1979),
Kawecki etal. (1978)
Mice protection from lethal infection
with HSV-2
host/target: Swiss mice
Challenge virus: herpes simplex virus
type 2 (HSV-2) [neutralization test]
RNA Kawecki etal. (1978)
Anti-inflammatory Angiotensin I-converting enzyme
inhibitory activity
Alkali extract Vunduk etal. (2015)
Strong inhibition of 3α-hydroxysteroid
dehydrogenase (3α-HSD), hyalu-
ronate lyase and weak inhibition of
cyclooxygenase-1 (COX-1)
[Enzyme-based assays: (3α-HSD)-
assay according to the method of
Penning; N-cetyl-N-trimethylammo-
nium bromide assay according to the
method of Ferrante; COX-1 assay]
Polyporenic acid C; (3α,12α,25S)-
12-hydroxy-3-(3-methoxy-1,3-
dioxopropoxy)-24-methylene-
lanost-8-en-26-oic acid;
(3α,12α,25S)-3-(acetyloxy)-12-hy-
droxy-24- methylene-lanost-8-en-
26-oic acid
Wangun etal. (2004)
Mice protection from ear edema induc-
tion by 12-O-tetradecanoylphorbol-
13-acetate (TPA)
Mice ear edema model Polyporenic acid A; polyporenic acid
C; (3α,12α,25S)-3-[(carboxyacetyl)
oxy]-12-hydroxy-24-methyl-
ene-lanost-8-en-26-oic acid;
(3α,12α,25S)-12-hydroxy-3-[[(3S)-
3-hydroxy-5-methoxy-3-methyl-
1,5-dioxopentyl]oxy]-24- meth-
ylene-lanost-8-en-26-oic acid;
(+)-12α,28-dihydroxy-3α-(30-h-
ydroxy-30-methylglutaryloxy)-24-
methyllanosta-8,24(31)-dien-26-oic
acid
Kamo etal. (2003)
Antioxidant Antioxidant capacity [DPPH scavenging activity, FRAP
method]
Water extracts Vunduk etal. (2015)
Antioxidant capacity [DPPH scavenging activity, reducing
power, α-carotene bleaching inhibi-
tion]
α-, β-, γ-, δ-tocopherols; ascorbic acid;
β-carotene; lycopene
Reis etal. (2011)
Antioxidant capacity [FRAP method] p-hydroxybenzoic acid; protocatechuic
acid; vanillic acid
Sułkowska-Ziaja etal. (2012)
Immunomodu-lation Activation of neutrophils to production
of reactive oxygen forms
Neutrophils from human peripheral
blood [LDCL method]
Water extracts from fruiting bodies
and mycelium
Shamtsyan etal. (2004)
Anticancer Antimigrative properties Cancer cell lines: A549, HT-29, T47D,
TE671 [wound assay]
Ethanol extracts Pleszczyńska etal. (2016), Zwolińska
(2004), Żyła etal. (2005)
Cancer cell line:TE671 [wound assay] Ether extracts Zwolińska (2004)

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Table 1 (continued)
Biological activity Mechanism of biological activity Model [method of study] ExtractaActive compoundaReferences
Cancer cell lines: A549, C6, HT-29,
T47D [wound assay]
Water extracts Pleszczyńska etal. (2016), Lemieszek
etal. (2009)
Cancer cell lines: A549, HT-29, T47D
[wound assay]
Water and ethanol extracts of culti-
vated fruiting bodies
Pleszczyńska etal. (2016)
Decrease in tumor cell adhesion Cancer cell line: LS180 [crystal violet
assay]
Ethanol and ether extracts of invitro
grown mycelium
Cyranka etal. (2011)
Apoptosis induction Cancer cell line: T47D [ELISA] Ethanol extracts Zwolińska (2004)
Cancer cell line: A549 [ELISA] Ether extracts Żyła (2005)
Cancer cell lines: A549, C6 [ELISA,
May Grünwald Giemsa staining]
Water extracts Lemieszek etal. (2009)
Cell death induction Cancer cell lines: A549, T47D, TE671
[May Grünwald Giemsa staining]
Ethanol extracts Żyła etal. (2005), Zwolińska (2004)
Decrease in cancer viability Cancer cell line: HeLa [MTT test] carboxymethylated (1→3)- -α-D-
glucans
Wiater etal. (2011)
Decrease in cancer viability Cancer cell line: LS180]MTT test] Ethanol and ether extracts of invitro
grown mycelium
Cyranka etal. (2011)
Inhibition of MMP-3, MMP-9,
MMP-14
Cancer cell line: A549 [zymography] Ethanol and ether extracts Zwolińska (2004)
Inhibition of MMP-9 Cancer cell line: HT-29 [zymography] Water extracts Lemieszek (2008)
Inhibition of MMP-1, MMP-3, MMP-9 [Hydrolysis of MMP protein sub-
strates—labeled synthetic peptides]
(E)-2-(4-hydroxy-3-methyl-2-butenyl)-
hydroquinone
Kawagishi etal. ( 2002)
Inhibition of MMP-1 [Hydrolysis of MMP protein sub-
strates—labeled synthetic peptides]
polyporenic acid C Kawagishi etal. (2002)
Inhibition of cancer cells proliferation Cancer cell lines: A549, C6, HEp-2,
HT-29, Jurkat E6.1, RPMI 8226,
T47D, TE671 [MTT test]
Ethanol extracts Pleszczyńska etal. (2016), Wasyl
(2006), Żyła etal. (2005), Zwolińska
(2004)
Cancer cell lines: A549, HT-29, T47D
[MTT test]
Ethanol extracts of cultivated fruiting
bodies
Pleszczyńska etal. (2016)
Cancer cell lines: A549, C6, FTC238,
HEp-2, HeLa, HT-29, Jurkat E6.1,
RPMI 8226, SK-N-AS, T47D,
TE671 [MTT test]
Ether extract Wasyl (2006), Kaczor etal. (2004),
Zwolińska (2004)
Cancer cell lines: A549, C6, HT-29,
Jurkat E6.1, T47D [MTT test]
Water extracts Pleszczyńska etal. (2016), Lemieszek
etal. (2009), Zwolińska (2004)
Cancer cell lines: A549, HT-29, T47D
[MTT test]
Water extracts of cultivated fruiting
bodies
Pleszczyńska etal. (2016)
Cancer cell lines: A549, T47D [MTT
test]
Polyporenic acid A Zwolińska (2004)
Inhibition of DNA synthesis Cancer cell line: C6 [BrdU test] Ethanol extracts Wasyl (2006)
Cancer cell lines: A549, C6 [BrdU
test]
Water extracts Lemieszek etal. (2009)
Alterations in cell cycle progression—
accumulation of cancer cells in the
“S” phase
Cancer cell line: FTC238 [flow
cytometry]
Ether extract Kaczor etal. (2004)
Inhibition of cancer cell growth Mouse sarcoma S-37 [not given] Extracts Blumenberg and Kessler (1963)

World J Microbiol Biotechnol (2017) 33:83
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resistance and therefore further increase the effective-
ness of the active substances. Particularly noteworthy
among the wide variety of biological activities of F.
betulina extract, are properties proved in invivo studies,
e.g. the efficacy of water and ethanol extracts in treat-
ment of the genital tract in dogs (Utzig and Samborski
1957; Wandokanty etal. 1954, 1955) or mice protection
from lethal infection with the TBE virus by water, etha-
nol, and ether extracts (Kandefer-Szerszeń et al. 1981;
Kandefer-Szerszeń and Kawecki 1974, 1979). The broad
spectrum of antiviral and antimicrobial activity of F. bet-
ulina extracts proved by a number of research teams in
different models based on different techniques deserves
special attention as well (see references cited in Table1).
Recently, Stamets (2011, 2014) has invented formulations
prepared from different medicinal mushrooms including
F. betulina, which are useful in preventing and treating
viral and bacterial diseases, i.e. herpes, influenza, SARS,
hepatitis, tuberculosis, and infections with E. coli and S.
aureus .
Some pure compounds corresponding to the bioactiv-
ity of the birch polypore were also identified (Fig.2).
They belong to several chemical classes but the greatest
attention was paid to small molecular weight second-
ary metabolites, especially triterpenoids. Kamo et al.
(2003) isolated several triterpenoid carboxylic acids
with a lanostane skeleton, e.g. polyporenic acids and
their derivatives (Table 1). In in vivo tests, the sub-
stances suppressed TPA-induced mouse ear inflamma-
tion up to 49–86% at the dose of 0.4µM/ear. Alresly etal.
(2016) purified one previously unknown (identified as
3β-acetoxy-16α hydroxyl-24-oxo-5α-lanosta-8-ene-21-
oic acid) and ten known triterpenes from ethyl acetate
extract of fruiting bodies of the fungus. The new com-
pound showed anti-gram-positive bacteria activity. The
medicinal activity of some triterpenoids tested was exam-
ined more accurately. It was shown that polyporenic acid
C, just like another compound isolated from F. betulina,
i.e. (E)-2-(4-hydroxy-3methyl-2-butenyl)-hydroquinone,
had inhibitory activity against some matrix metallopro-
teinases (MMP), with IC50 values (concentration causing
inhibition by 50% compared to control) in the range from
23 to 128µM (Kawagishi etal. 2002). Polyporenic acid C
and three other F. betulina triterpenoids (Table1) showed
anti-inflammatory and antibacterial activity by strong
inhibition of 3α-hydroxysteroid dehydrogenase and bac-
terial hyaluronate lyase activity, respectively (Wangun
etal. 2004).
In their search for fungal antimicrobial substances,
Schlegel et al. (2000) isolated another valuable com-
pound—piptamine, N-benzyl-N-methylpentadecan-
1-amine from submerged culture of F. betulina Lu 9-1.
It showed activity against gram-positive bacteria (MIC,
Table 1 (continued)
Biological activity Mechanism of biological activity Model [method of study] ExtractaActive compoundaReferences
Tumor size reduction by induction of
cancer cell necrolysis and inhibition
of metastases
Female dogs with adenocarcinoma
mammae, adenocarcinoma solidum,
adenocarcinoma papilliferum
[histopathological examination after
Hansen staining]
Water extracts Pentacyclic triterpenes (suggested) Wandokanty etal. (1954; 1955)
Tumor size reduction and inhibition of
bleeding from the genital tract
Female dogs with Sticker’s sarcoma
[per vaginal examination]
Ethanol extracts Pentacyclic triterpenes (suggested) Utzig and Samborski (1957)
Neuroprotec-tion Protection of neurons against damage
induced by cisplatine, trophic stress,
excitotoxicity
Mouse neurons—10-day old [LDH
test]
Ethanol and ether extracts Wasyl (2006)
Cancer cell lines: A549—human Caucasian lung carcinoma, C6—rat glioma, FTC238—human thyroid carcinoma, HeLa—human cervical adenocarcinoma, Hep-2 (HeLa derivative)—human
cervix carcinoma, HT-29—human colon adenocarcinoma, Jurkat E6.1—human T-cell leukemia, LS180—human colorectal adenocarcinoma, RPMI 8226—human multiple myeloma, SK-N-
AS—human neuroblastoma, T47D—human breast ductal carcinoma, T671—human rhabdomyosarcoma/medulloblastoma
a Extracts/compounds were isolated from fruiting bodies of wild growing F. betulina, unless otherwise indicated
3α-HSD 3-α hydroxysteroid dehydrogenase, BrdU − 5-bromo-2’-deoxyuridine, COX-1 cyclooxygenase-1, DPPH 2,2-diphenyl-1-picrylhydrazyl, ELISA enzyme-linked immunosorbent assay,
FRAP ferric ion reducing antioxidant power, LDCL luminol-dependent chemiluminescence, LDH lactate dehydrogenase, MTT methylthiazolyldiphenyl-tetrazolium bromide, NCCLS National
Committee for Clinical Laboratory Standards

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83 Page 8 of 12
Fig. 2 Chemical structures of bioactive compounds isolated from F. betulina

World J Microbiol Biotechnol (2017) 33:83
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Page 9 of 12 83
minimum inhibitory concentration, values in the range
from 0.78 to 12.5 µg/ml) and yeasts including Candida
albicans (MIC 6.25µg/ml).
Polysaccharides from higher basidiomycota mushrooms
have been usually considered to be the major contributors
of their bioactivity. However, birch polypore polysaccha-
rides have not yet been sufficiently explored, in terms of
either the structure or pharmacological activity. It is known
that the Fomitopsis cell wall contains (1→3)-β-d-glucans
in an amount of ca. 52% dw (Jelsma and Kreger 1978;
Grün 2003). They are built from β-d-glucopyranose units
connected with (1→3)-linkages in the main chain, with
(1→3)-β-d linked side branches. However, there are no
reports about their biological activities. Another polysac-
charide isolated from the birch polypore was water-insolu-
ble, alkali-soluble (1→3)-α-d-glucan. Although α-glucans
are believed to be biologically inactive, its carboxymethyl-
ated derivative showed moderate cytotoxic effects invitro
(Wiater etal. 2011).
Miscellaneous applications
With the knowledge of the mechanisms of action of brown
rot decay, there are possibilities of new applications of
these fungi in biotechnology. The enzymatic and non-enzy-
matic apparatus for lignocellulose degradation can be used
for bioprocessing of biomass towards fuels and chemicals
(Arantes et al. 2012; Giles and Parrow 2011; Ray et al.
2010). Brown rot fungi, including F. betulina, were tested
for bioleaching of heavy metals (Cu, Cr, and As) from
wood preservatives due to accumulation of metal-complex-
ing oxalic acid (Sierra Alvarez 2007). Production of bio-
mass degrading enzymes, for instance cellulases, hemicel-
lulases, amylases, etc., was also studied (Krupodorova etal.
2014; Valášková and Baldrian 2006a, b).
The cell wall of F. betulina can be a source of useful pol-
ysaccharides, e.g. water-insoluble, alkali-soluble α-glucans
(Grün 2003; Jelsma and Kreger 1979). (1→3)-α-d-glucans
whose main chain contains 84.6% of (1→3)-linked α-d-
glucopyranose in addition to 6% of (1→4)-linked units were
purified and characterized by Wiater etal. (2011). Another
polysaccharide, named piptoporane I, was extracted and
purified by Olennikov etal. (2012). This α-glucan was built
from residues of (1→3)-α-d-glucopyranose with occasional
branching by single residues of β-d-glucopyranose at the C6
position (17.3%). It has been shown that fungal (1→3)-α-d-
glucans, including that from F. betulina, effectively induce
the production of microbial (1→3)-α-glucanases (mutan-
ases), i.e. enzymes that have potential in dental caries
prevention. (1→3),(1→6)-α-d-Glucans (mutans) synthe-
sized by mutans streptococci are key structural and func-
tional constituents of dental plaque matrix; therefore, they
seem to be a good target for enzymatic anti-caries strategy
(Pleszczyńska etal. 2015). However, streptococcal glucans
are difficult to use as inducers of mutanases because of the
low yield and structural variation. Birch polypore α-glucan,
whose amount in the cell wall of F. betulina reaches even
44–53% dw (Grün 2003), can be used to replace strepto-
coccal glucans (Wiater etal. 2008).
Conclusions andoutlook
The F. betulina fungus has been widely used and appreci-
ated in folk medicine, and modern pharmacological studies
have confirmed its potential indicating significant antimi-
crobial, anticancer, anti-inflammatory, and neuroprotective
activities. The possibility of successful cultivation thereof
in artificial conditions additionally promotes the applica-
bility of the fungus. However, compared with other poly-
pore fungi, the research on F. betulina is less developed;
for instance, little is known about its lifestyle, including
the wood degradation strategy. Moreover, most of the bio-
activity studies have been performed using crude extracts;
hence, only a few of the effects have been associated with
the active substances identified, e.g. antibacterial activities
with piptamine or polyporenic acids. With a few excep-
tions, we still do not know the mechanisms underlying the
biological activities. Verification of biological activities
in invivo and clinical studies is also required. The further
research could contribute to better exploitation of the F.
betulina application potential.
Compliance with ethical standards
Competing interests The authors have no conflict of interest to
declare.
Open Access This article is distributed under the terms of the
Creative Commons Attribution 4.0 International License (http://
creativecommons.org/licenses/by/4.0/), which permits unrestricted
use, distribution, and reproduction in any medium, provided you give
appropriate credit to the original author(s) and the source, provide a
link to the Creative Commons license, and indicate if changes were
made.
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