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Effects of Agaricus brasiliensis mushroom in
Walker-256 tumor-bearing rats
Fernanda Menon Dias Jumes, Daiana Lugarini, Amanda Leite Bastos Pereira,
Anabel de Oliveira, Adriana de Oliveira Christoff, Giani Andrea Linde,
Juliana Silveira do Valle, Nelson Barros Colauto, and Alexandra Acco
Abstract: Agaricus brasiliensis is a mushroom native to Sa
˜o Paulo State, Brazil, that is studied for its medicinal propri-
eties. This work aimed to investigate the antitumoral activity of A. brasiliensis extracts and pure powdered basidiocarp
preparation using Walker-256 (W256) tumor-bearing rats, a model for cancer-related cachexia studies. The rats were
treated for 14 days by gavage (136 mg/kg) and at the end of the experiment tumors were collected to calculate mass and
volume. Blood was collected for determination of plasma glucose, albumin, alanine aminotransferase (ALT), and aspartate
aminotransferase (AST). Hepatic and tumor enzymes indicating oxidative stress were also evaluated. The results showed
that all 4 treatments (pure powdered basidiocarp and aqueous, acid, and alkaline extracts) significantly reduced tumor size
and promoted gain in body weight. Plasmatic analysis showed a reduction in AST level and increased glycemia in the
treated rats. Pure basidiocarp preparations improved the liver catalase and superoxide dismutase activity, but did not
change the glutathione S-transferase activity. The data collected from the W256 tumor-bearing rats revealed the beneficial
effects of A. brasiliensis in tumor treatment, mainly related to cachexia. The benefits can be partly related to antioxidant
activity and to reduction of weight loss and tumor growth.
Key words: Agaricus brasiliensis,Agaricus blazei, Walker-256, tumor, liver, oxidative stress.
Re
´sume
´:Agaricus brasiliensis est un champignon originaire de Sa
˜o Paulo, au Bre
´sil, e
´tudie
´pour ses proprie
´te
´sme
´dicina-
les. La pre
´sente e
´tude a pour but d’examiner l’activite
´antitumorale d’extraits de A. brasiliensis et d’une pre
´paration de ba-
sidiocarpe en poudre pur, en utilisant des rats porteurs de la tumeur Walker-256 (W256), un mode
`le pour les e
´tudes de la
cachexie lie
´e au cancer. On a traite
´les rats pendant 14 jours par gavage (136 mg/kg) et extrait la tumeur a
`la fin de l’ex-
pe
´rience pour calculer la masse et le volume. On a pre
´leve
´le sang pour de
´terminer le glucose sanguin, l’albumine,
l’alanine aminotransfe
´rase (ALT) et l’aspartate aminotransfe
´rase (AST). On a aussi e
´value
´les enzymes he
´patiques et tumo-
rales indicatrices de stress oxydatif. Les re
´sultats ont montre
´que quatre traitements complets (extraits de basidiocarpe en
poudre pur, aqueux, acide et alcalin) ont re
´duit significativement la taille de la tumeur et favorise
´le gain de poids corpo-
rel. L’analyse plasmatique a montre
´une re
´duction du taux d’AST et une augmentation de la glyce
´mie chez les rats traite
´s.
La pre
´paration de basidiocarpe pur a ame
´liore
´les taux de catalase et de superoxyde dismutase he
´patiques, mais n’a pas
modifie
´les taux de glutathion-S-transfe
´rase. Les donne
´es obtenues chez les rats porteurs de la tumeur W-256 ont de
´montre
´
les effets be
´ne
´fiques de A. brasiliensis dans le traitement des tumeurs principalement associe
´es a
`la cachexie. Les effets
be
´ne
´fiques peuvent e
ˆtre lie
´s en partie a
`l’activite
´antioxydante, re
´duisant la perte de poids et la grosseur des tumeurs.
Mots-cle
´s : Agaricus brasiliensis,Agaricus blazei, tumeur Walker-256, tumeur, foie, stress oxydatif.
[Traduit par la Re
´daction]
Introduction
Agaricus brasiliensis Wasser (formerly A. blazei Murrill
(sensu Heinemann) according to Wasser et al. (2002), and
considered by Kerrigan (2005) as A. subrufescens Peck) is a
mushroom native to Sa
˜o Paulo State, Brazil. Many biologi-
cal actions are attributed to this fungus, such as immunomo-
dulation (Ito et al. 1997; Ebina and Fujimiya 1998; Fujimiya
et al. 1999; Sorimachi et al. 2001; Kaneno et al. 2004; Kasai
et al. 2004), protection of genetic damage (Delmanto et al.
2001; Martins de Oliveira et al. 2002; Luiz et al. 2003; Bar-
bisan et al. 2003; Machado et al. 2005; Bellini et al. 2006),
inhibition of tumor cell growth, and inhibition of cell migra-
tion or tumor-induced neovascularization (Kawamura and
Kasai 2007; Takaku et al. 2001).
The therapeutic effects of A. brasiliensis are attributed to
b-D-glucans, which are polysaccharides present in the cell
wall that can bind to proteins (Mizuno et al. 1999). The b-
glucan receptor has been identified as a b-glucan inhabitable
receptor for particulate activators of the alternative comple-
ment pathway (Czop and Austen 1985), such as complement
receptor type 3 (CR3; also known as Mac-1, CD11b/CD18,
or aMb2-integrin), which is the major b-glucan receptor on
Received 28 May 2009. Accepted 15 September 2009. Published
on the NRC Research Press Web site at cjpp.nrc.ca on
14 January 2010.
F.M.D. Jumes, D. Lugarini, A.L. Bastos-Pereira,
A. de Oliveira, A.O. Christoff, and A. Acco.1Pharmacology
Department, Federal University of Parana
´, Jardim das Ame
´ricas,
C.P. 19031, 81531-900 Curitiba –PR, Brazil.
G.A. Linde, J.S. doValle, and N.B. Colauto. Molecular
Biology Laboratory, Paranaense University, Prac¸a Mascarenhas
de Moraes, 4282, C.P. 224, 87502-210 Umuarama – PR, Brazil.
1Corresponding author (e-mail: aleacco@ufpr.br).
21
Can. J. Physiol. Pharmacol. 88: 21–27 (2010) doi:10.1139/Y09-111 Published by NRC Research Press
leukocytes (Ross et al. 1999; Xia et al. 1999). More re-
cently, another receptor, dectin-1, has also been identified
as a b-glucan receptor (Brown et al. 2003). Beyond its ac-
tion in specific receptors, extracts of A. brasiliensis have
also shown in vitro antioxidant activity in the DPPH (2,2-
diphenyl-1-picrylhydrazyl) assay (Oliveira et al. 2007;
Nakajima et al. 2007; Watanabe et al. 2008), but this antiox-
idant action still requires more study in vivo, particularly in
the enzymes involved in the oxidative stress process. As re-
active oxygen (ROS) and nitrogen (RNS) species play a sig-
nificant role in cancer pathogenesis (Lev et al. 2003),
antioxidant compounds can be therapeutically useful in neo-
plasias. Many medicinal mushrooms contain polyphenols,
commonly termed free radical-scavenging molecules (Lee et
al. 2007). However, the biological activity of b-D-glucans is
influenced by their solubility in water (Ishibashi et al. 2001),
molecular weight (Okazaki et al. 1995; Cleary et al. 1999;
Mueller et al. 2000), branching rate (Cleary et al. 1999; Ka-
taoka et al. 2002), triple helical solution conformation (Mu-
eller et al. 2000; Falch et al. 2000), and b-(1?6)-bonding
system in the b-(1?3) major chain (Cleary et al. 1999). In
this sense, the pH used in the extraction process could per-
haps affect the b-D-glucan chains and activity.
Various cancer models have been reportedly used to test
the antitumoral activity of A. brasiliensis, in vivo and in vi-
tro, such as sarcoma 180 in mice (Ohno et al. 2001; Lee et
al. 2003; Fan et al. 2007), Ehrlich’s tumor in mice (Kaneno
et al. 2004), V79 cells in hamsters (Menoli et al. 2001), hu-
man breast cancer cells (MCF7) (Talorete et al. 2002), and
Lewis cells of lung carcinoma in mice (Takaku et al. 2001).
However, no reports were found on tumor Walker-256
(W256). This fast-growing carcinosarcoma is an appropriate
model for studying the cachexia syndrome and anticancer
treatments in rats because it is species-specific and easily
transplanted (Guaitani et al. 1982). Within a short time after
its inoculation, there is a reduction in body weight, anorexia,
and difficulty in the catabolism of proteins, carbohydrates,
and lipids. In 14 days, the tumor mass may represent a con-
siderable percentage of rat body weight with potential occur-
rence of death (Vicentino et al. 2002).
This study aimed to evaluate the influence of A. brasilien-
sis extracts in rats bearing the W256 tumor as a cancer
model to verify the antitumoral activity and physiological
parameters related to hepatic function and oxidative stress.
Materials and methods
Production and extracts of A. brasiliensis
The extracts were prepared with basidiocarps of ABL97/
11 A. brasiliensis strain from the Molecular Biology Labora-
tory at Paranaense University (Umuarama, Brazil). Com-
posting method and raw materials were used to produce
basidiocarps according to Colauto et al. (2010). The basidio-
carps were harvested, washed with water, dehydrated for
48 h at 65 8C, powdered, and stored in polypropylene bags
at –70 8C. The A. brasiliensis extracts were prepared with
powdered basidiocarp mixed 1:10 (m/v) with ultra pure
water only or pH-adjusted to 4.0 with HCl (1.0 mol/L) or to
8.0 with NaOH (1.0 mol/L). Mixtures were then kept in a
closed glass flask at 90 8C for 16 h (Souza-Paccola et al.
2004). Later, all extracts were filtered in a vacuum system,
using paper filter number 11 (Qualy) and had the final pH
adjusted to 5.4. All mushroom extracts were divided into in-
dividual aliquots and stored at –20 8C. Daily aliquots were
defrosted at room temperature to be administered to rats.
Tumor W256 implantation
W256 carcinoma cells were suspended in phosphate-
buffered saline (16.5 mmol/L phosphate, 137 mmol/L NaCl,
2.7 mmol/L KCl, pH 7.4) and injected into the right flank of
the rats on day 1 of the experiment. The W256 cells were
injected subcutaneously (2 107cells) after cell viability as-
sessment by the trypan blue exclusion test in a Neubauer
chamber. Maintenance of W256 cells was carried out by
weekly intraperitoneal (i.p.) inoculation, according to Vicen-
tino et al. (2002).
Treatments and biological material
Male Wistar rats (260 ± 30 g) were obtained from the
Central Animal House of the Federal University of Parana
´
(Curitiba, Brazil). The rats were placed in a room at
22 8C±18C under a 12 h light : 12 h dark cycle with free
access to standard laboratory food (Purina) and tap water.
The Institutional Animal Ethics Committee (CEEA), which
works in accordance with the Canadian Council on Animal
Care, approved the procedures, and the project received cer-
tificate number 174.
To assess the biochemistry data as well as body and tu-
mor mass, the tumor-bearing rats were divided into 5 treat-
ment groups, namely, (i) control (n= 5), saline solution;
(ii) PureB (n= 6), pure powdered basidiocarp added to sal-
ine solution in a proportion of 1:10; (iii) Eaque (n= 6),
aqueous extract of the mushroom powder; (iv) Eacid (n=
6), acid extract of the mushroom powder; and (v) Ealka
(n= 6), alkaline extract of the mushroom powder. The treat-
ment started 1 day after cell implant, and continued for
14 days, once a day by gavage, with a daily dose of
136 mg/kg. This dose was calculated by interspecific allo-
metric scaling (Pachaly and Brito, 2000) on the basis of
doses previously published for mice (Lee et al., 2003; Ka-
neno et al. 2004). Every week and at the end of the experi-
ment, the body mass of the rats was evaluated. On the last
day of treatment, the rats were kept on a 12 h fast and then
anesthetized with an i.p. injection of thiopental (70 mg/kg).
The blood was collected with a heparin-syringe from the ab-
dominal cava vein and centrifuged to separate the plasma
(3000gfor 10 min). Finally, euthanasia was induced. The tu-
mor was collected and weighed in an analytical balance.
Both the tumoral volume and the inhibition growth rate
were calculated by measuring the tumor diameters, accord-
ing to the study of Mizuno et al. (1999).
Biochemistry
After assessing the plasma on the last day of the experi-
ment, biochemical analysis was conducted for glucose, albu-
min, alanine aminotransferase (ALT), and aspartate
aminotransferase (AST) utilizing commercial kits from
Labtest Diagnostica (Lagoa Santa, Brazil).
Oxidative stress
To assess the oxidative stress enzyme activities in both
liver and tumor, a slightly different treatment was used. The
22 Can. J. Physiol. Pharmacol. Vol. 88, 2010
Published by NRC Research Press
tumor-bearing rats were separated into 2 groups, control
(n= 5) and PureB (n= 6), and both were treated for
14 days as described above. In addition, a 3rd group was
added, namely basal (n= 5), which included only healthy
untreated rats without tumor. These animals were used as
well for determination of body mass. On the 15th day, frac-
tions of liver and tumor were collected and stored at –70 8C
for further determination of catalase (CAT), superoxide dis-
mutase (SOD), and glutathione S-transferase (GST) activ-
ities. Liver and tumor samples were homogenized in
phosphate buffer (pH 6.5) for enzyme analysis. Catalase ac-
tivity was measured following the method of Aebi (1984).
SOD activity was measured by the ability of this enzyme to
inhibit pyrogallol autoxidation at 440 nm (Gao et al. 1998).
The amount of enzyme that inhibited the reaction by 50%
(IC50) was defined as one unit of SOD, and the enzyme ac-
tivity was expressed in units of SOD per milligram of total
protein (U/mg). GST activity was measured by the proce-
dures of Habig et al. (1974). Total protein concentration in
tissue homogenates was determined by the Bradford method
(Bradford 1976).
Statistical analysis
Mean values ± standard error of the mean (SE) were cal-
culated. The data were analyzed statistically by variance
analysis (ANOVA) and Tukey test for comparison of aver-
ages. Values were considered significant when p< 0.05.
Results and discussion
The present data showed the beneficial effect of the
A. brasiliensis extracts in cachexia related with cancer. The
variation in the body mass, represented by the difference in
the body weight from the first and the last day of the experi-
ment and discounting the tumor mass, is presented in Table
1. All the rats that received the A. brasiliensis extracts and
pure (raw) powdered basidiocarp (PureB) increased body
mass throughout the treatment, unlike the control group
(–14.17 ± 24.8 g). That result was expected for the control
rats as the W256 tumor is a model that correlates cachexia
and cancer, its main characteristic being damage to or less-
ening of body weight. Increase in body mass was less evi-
dent in the PureB group (32.65 ± 8.9 g). The highest
increase in body mass occurred in the Ealka group (54.69 ±
7.9 g), which was similar to the gain weight presented by
the basal group (58.65 ± 4.86 g) composed of untreated non-
tumoral rats. These results reveal that all the A. brasiliensis
extracts and pure powdered basidiocarp tested were able to
protect the rats against loss of body weight induced by tu-
mor.
According to the inhibition growth rate (IGR), as shown
in Table 1, all the extracts and powdered basidiocarp signif-
icantly reduced the tumor growth when compared with the
control group. The greatest inhibition (92.02%) occurred in
the rats treated with pure extract. Both aqueous and alkaline
extracts evidenced similar inhibition rates, while the lowest
rate, although still high, was observed in the group treated
with acid extract (74.18%). Although complete tumor sup-
pression was not found in any treated group, the IGR was
pronounced for all the mushroom preparations, which were
extracted using different pH conditions. Apart from the large
commercial and clinical use of A. brasiliensis around the
world, these data indicate that the products or extracts
should be properly prepared to actively maintain the com-
pounds responsible for antitumoral effects.
The weight loss in cancer, as that observed in the control
group, is associated with anorexia, which is a spontaneous
loss of appetite after cachexia, characterized by severe innu-
trition, intense muscular and adipose catabolism, anemia,
negative nitrogen balance, and metabolic alterations (Silva
2006). Glucose and protein metabolism is affected, showing
variation. Our data revealed that the treatments did not
change the albumin level (Fig. 1A), but the mean values in
all groups fell below the reference index for rats (4.14 ±
0.21 g/dL; Kaneko 1989). However, Gu et al. (2005) ob-
served a significant difference in the albumin level 18 days
after inoculation of W256 cells directly into rat livers. The
present experiment was conducted for 14 days and finalized
before that period. The lack of significance in our data can
also be correlated with the albumin half-life, which is
21 days (Cardillo-Vieira et al. 1995).
Glucose metabolism is frequently modified to meet the
nutritional exigencies in illnesses like neoplasia. Cancer
cells use glucose as the main substrate, 10 to 50 times more
than normal cells (Guppy et al. 2002). Despite an expected
decrease in the glucose plasmatic level in this situation,
sometimes it does not occur because of an increase in the
amino acids and lactate hepatic gluconeogenesis (Silva
2006). When comparing the plasmatic glucose level in all
the treated groups, there was no significant difference (Fig.
Table 1. Effect of treatment with Agaricus brasiliensis preparations on body mass, tumor volume and mass, and in-
hibition growth rate in Walker-256 tumor-bearing rats after 15 days of carcinoma cell implantation.
Treatment groups Body mass variation, g Tumor mass, g Tumor volume, cm3Inhibition growth rate, %
Control –14.17 ± 24.8 28.08 ± 10.5 206.50 ± 71.7 —
PureB 32.65 ± 8.9 3.56 ± 1.2* 16.47 ± 6.4** 92.02
Eaque 49.85 ± 6.8 7.37 ± 3.3* 30.68 ± 15.7** 85.14
Eacid 44.57 ± 7.6 7.87 ± 1.8* 53.30 ± 10.7* 74.18
Ealka 54.69 ± 7.9* 4.90 ± 1.5* 31.26 ± 12.0** 84.88
Basal 58.65 ± 4.8** — — —
Note: Wistar rats were treated as described in Materials and methods. Treatment groups: Control, saline solution; PureB, pure
powdered basidiocarp; Eaque, aqueous extract; Eacid, acid extract (pH 4.0); Ealka, alkaline extract (pH 8.0); and Basal, untreated,
non-tumor-bearing rats. Body mass variation was calculated by the difference in mass between the 15th and the 1st experimental day,
discounting the tumor mass. Tumor volume, V(cm3), was calculated as (4p/3a2)(b/2), where ais the minor diameter and bis the
major diameter. Inhibition growth rate (%) was defined as (1 – T/C) 100, where T is treated and C is control. *, Significant at p<
0.05 and **, p< 0.01 compared with control (ANOVA and Tukey test).
Jumes et al. 23
Published by NRC Research Press
1B). However, there was a strong tendency to increase the
glycemia in the PureB, Eaque, and Ealka groups, and these
treatment groups had mean values comparable with the
reference levels for rats (47.70 to 107 mg/dL; Kaneko
1989). The increase observed was 147%, 103%, and 142%
higher than control in the rats treated with pure powdered
basidiocarp, aqueous extract, and alkaline extract, respec-
tively. This glycemia elevation could be a direct result of
the A. brasiliensis intake because 55% of its composition
can be carbohydrates, including the active b-glucan com-
pounds (Firenzuoli et al. 2008), which are polysaccharides
that contain only glucose as structural components. The ab-
sorption of glucans after oral adminstration was previosly
reported in mice (Suzuki et al. 1991). However, the mecha-
nism by which large b-glucans could be taken up by the gas-
trointestinal tract still remains unknown (Firenzuoli et al.
2008).
A lower glycemia level occurred in the Eacid group
(44.93 ± 10.23 mg/dL) despite the fact that those rats had
exhibited increased body mass during the 15 days of treat-
ment. These data reveal that the glycemia is only one factor
that determines the change in body condition. Other impor-
tant metabolic abnormalities can also induce body mass var-
iations in cancer patients (Silva 2006), such as alterations in
the hepatic metabolism, which is profoundly affected (Ka-
zantzis and Seelaender 2005). Regarding liver enzyme meas-
urements, ALT remained at the same level in all the groups.
However, AST levels dropped in the treated groups when
compared with the control (Fig. 1C). These data indicate
that tumor-induced alterations in the permeability or func-
tion of the hepatocytes resulted in extravasation of AST for
the plasma in the control rats. It may occur because of the
AST cytosolic localization in hepatocytes (Jagadeesan and
Kavitha 2006). Our data indicate that A. brasiliensis ex-
tracts, partially at least, protected the hepatocytes, resulting
in a reduced level of plasmatic AST. This cellular protection
may be due to higher CAT and SOD activity, both enzymes
that reduce the action of free radicals in proteins, enzymes,
and membranes (Gate
´et al. 1999; Deweese et al. 2001).
When both hepatic and tumoral tissues were compared,
higher catalase activity was observed in the liver, mainly in
the basal group (Table 2). The presence of the W256 tumor
significantly reduced the hepatic enzyme activity, which
corroborates the literature (Bastos-Pereira et al. 2009).
Some tumors have reduced capacity to detoxify hydrogen
peroxide (H2O2) because of a lower CAT level (Valko et al.
2006). The treatment with A. brasiliensis raw powdered ba-
sidiocarp increased by 2-fold the hepatic activity of CAT
(123.7 ± 15.37 mmolmin–1mg–1 protein) compared with
control rats (60.76 ± 14.32 mmolmin–1mg–1 protein). These
data reveal the antioxidant effect of the basidiocarp, as the
CAT efficiently promotes the conversion of H2O2to water
Fig. 1. Plasmatic levels of (A) albumin, (B) glucose, (C) aspartate aminotransferase (AST), and (D) alanine aminotransferase (ALT) in
Walker-256 tumor-bearing rats treated for 14 days with Agaricus brasiliensis mushroom preparations (pure, aqueous, acid, or alkaline) or
saline solution (control) by gavage. Values are means ± SE. *, Significant at p< 0.05 and **, p< 0.01 compared with control (ANOVA and
Tukey test).
24 Can. J. Physiol. Pharmacol. Vol. 88, 2010
Published by NRC Research Press
and molecular oxygen. In the tumor, only a discrete but not
significant increase in the CAT activity was observed.
In the same way as the CAT enzyme, hepatic SOD activ-
ity was reduced in the presence of tumor in the control
group (1.58 ± 0.04 U/mg protein). The treatment signifi-
cantly elevated the hepatic SOD (1.96 ± 0.14 U/mg protein),
raising it to the basal levels (1.99 ± 0.16 U/mg protein)
(Table 2). The levels of tumoral SOD in both groups were
increased approximately 2.7-fold when compared with the
hepatic enzyme. This stimulus occurs, at least in part, be-
cause tumors produce an oxidant condition in many tissues,
which may increase the SOD activity attending to attenuate
superoxide radical (O2–). This primary ROS reacts with
other molecules inducing secondary ROS. The treatment
with A. brasiliensis, however, did not reduce the tumoral
SOD. As hepatic SOD is the first-line defense against oxida-
tive stress (Gate
´et al. 1999), this indicates that W256 tumor
is a high-oxidant tissue. A positive correlation between SOD
activity and the phenol compounds present in medicinal
mushrooms was earlier demonstrated (Kim et al. 2008). In
this context, A. brasiliensis could be considered a rich
source of compounds that promote its antioxidant activity.
GST is the other enzyme that plays a role in cell defense
against many endogenous and environmental carcinogens by
the conjugation of glutathione (GSH) with reactive or oxi-
dant electrons (Deweese et al. 2001). However, it seems
that hepatic GST is not involved in the A. brasiliensis ef-
fects, because no significant difference was found between
the treated and control groups, despite both groups showing
a significantly reduced level when compared with the basal
group (Table 2). GST activity in the W256 tumor could not
be measured, probably because of its very low expression in
this tissue. High expression can be expected in tissues with
elevated biotransformation rate, such as the liver.
Various extracts and isolated compounds from
A. brasiliensis have been tested in studies with cancer cell
lines and animal cancer models under different protocols of
dose, administration, and time of therapy. The most-cited
solid tumor models are sarcoma 180 (Lee et al. 2003; Fan
et al. 2007; Liu et al. 2008; Moura
˜o et al. 2009) and Ehrlich
adenocarcinoma (Ito et al. 1997; Kaneno et al. 2004; Pinto
et al. 2009), both in mice. Most of the articles have shown
antitumoral and immunomodulating activity, despite the de-
scription of different action mechanisms in these biological
processes. Although rat tumor models have been less fre-
quently used in studies involving A. brasiliensis, the present
data reinforce the beneficial effects of extracts of this mush-
room and pure powdered basidiocarp in the treatment of
solid tumors.
This is the first report showing the effects of
A. brasiliensis in cachexia studied with W256 tumor. The
benefits may be partly related to antioxidant activity, beyond
reduction in body mass loss and tumor growth. More studies
should be encouraged to identify the cellular protection
mechanisms in cachexia and the effects after prolonged ther-
apeutic use, aspects that were not evaluated in the present
study.
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Table 2. Enzymatic activity of CAT, SOD, and GST in liver and tumor homogenate obtained after 15 days of
subcutaneous inoculation of Walker-256 tumor cells in rats.
Basal Control PureB
Enzyme Liver Liver Tumor Liver Tumor
CAT, mmolmin–1mg–1 460.8±37.58* 60.76±14.32 36.5±1.67 123.7±15.37 41.39±3.46
SOD, Umg–1 1.99±0.16 1.58±0.04* 4.43±0.26 1.96±0.14 5.36±0.58
GST, pmolmin–1mg–1 607.5±29.52** 379.9±69.75 — 357.0±19.56 —
Note: CAT, catalase; SOD, superoxide dismutase; GST, glutathione S-transferase. Control and pure basidiocarp (PureB)
groups were conducted with rats inoculated with W256 cells, as described in Materials and methods, and treated with saline
solution or Agaricus brasiliensis powdered preparation, respectively. Rats in the basal group were neither inoculated with tu-
mor cells nor treated. *, Significant at p< 0.05 and **, p< 0.01 for each enzyme compared with the other groups in the same
tissue (liver or tumor) by ANOVA and Tukey test.
Jumes et al. 25
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