Original Articles

Anti-Herpes Simplex Virus Effect of an Aqueous Extract of Propolis

Mahmoud Huleihel PhD and Vladimir Isanu MD

Institute for Applied Biosciences, Ben-Gurion University of the Negev, Beer Sheva, Israel

Key words: propolis extract, herpes simplex virus, antiviral activity, plaque assay, cytopathic effect

Abstract

Propolis, a natural product from beehives, comprises a complex of chemicals, the most important group being
flavinoids, which play a role in antiviral protection.
To test the inhibitory effect of propolis extract against
herpes simplex viruses in vitro and in vivo.
propolis was added to Vero cells at various
times and concentrations before, at or after infection with HSV-1. In
vivo the effect of propolis was tested in newborn rats infected s.c. or
i.p. and on rabbit corneas infected with HSV-1.
In vitro 0.5% propolis extract caused 50% inhibition of
HSV infection. There was indirect evidence for a strong interaction
between the propolis extract and the surface of the Vero cells, but there
was no direct interaction with HSV-1 particles. Administration of
propolis before or at the time of infection yielded the most significant
inhibitory effect, but even when 10% propolis extract was added 2
hours post-infection it gave 8085% protection. In vivo as little as 5%
propolis prevented the appearance and development of symptoms of
local and i.p. HSV-1 infection in rats and of corneal HSV-1 infection in
rabbits. There were no cytotoxic effects at a concentration of 10% in
vitro or 20% in vivo.
The potent antiviral activity of propolis against HSV1 infection in vitro and In vivo is probably due to prevention of virus
absorption into the host cells and/or inhibition of an internal step(s)
during the viral replication cycle.
Background:

Objectives:

Methods: In vitro:

Results:

Conclusions:

IMAJ 2002;4(Suppl):923927

Increasing efforts are currently being devoted towards finding

applications for natural products and their derivatives in the
treatment of human viral diseases. One such product is propolis.
This material, which is obtained from beehives, is based on resins
collected by bees from certain trees and plants. The chemical
composition of propolis is complex and has not been completely
elucidated. Nevertheless, it is known that the most important group
of compounds in terms of amount and biochemical activity is
the flavonoids, which are thought to play a significant role in the
antiviral protection process [14]. Over the past two decades the
material has been the subject of various experimental and clinical
studies investigating its antimicrobial, antifungal, anti-inflammatory, anti-tumor and anti-apoptotic properties [512]. A number of
authors have reported that propolis extracts have an inhibitory
effect on the development of an infection process caused by viruses
affecting plants (cucumber mosaic, tobacco spot, tobacco necrosis),
animals (HSV-1, varicella zoster and influenza), and humans
(human immunodeficiency). These findings have indicated the
potential of propolis as a possible antiviral drug [1,3,1317].
HSV = herpex simplex virus

IMAJ . Vol 4, Supplement . November 2002

In this work, we tested the inhibitory effects against HSV of an

aqueous extract of propolis in vitro on Vero cell cultures and in vivo
in rats and rabbits inoculated with the virus.
Materials and Methods
Cells and viruses

African green monkey kidney (Vero) cells were purchased from the
American Type Culture Collection, Rockville, MD, USA. Cells were
grown in RPMI medium containing 10% fetal calf serum, 1%
glutamine, antibiotics and incubated at 378C in humidified air
containing 5% CO2. HSV-1 and HSV-2 were also obtained from the
American Type Culture Collection. HSV-1 stock (3.106 pfu/ml) was
used for infecting the cells and the animals. HSV-2 stock (1.106 pfu/
ml) was used only for infecting cells.
Animals

The experiments were carried out on white, newborn (23 day old)
rats and young male (1.52 kg) rabbits. All animal experiments were
conducted in accordance with the ethical guidelines of the
Committee for the Ethical Care and Use of Laboratory Animals of
Ben-Gurion University of the Negev.
Propolis

Propolis was obtained from the beehives of Kibbutz Yad Mordekhai,

Israel. An aqueous extract of propolis (20% of raw propolis) was
prepared according to patent no. 101730 [18] and then diluted with
RPMI culture medium to give the required concentrations (0.1, 1, 5,
10 or 20%). The pH of the diluted extract was adjusted to 7.17.2
with sodium borate. As a control, we used phosphate-buffered
saline (pH 7.2) containing the same amount of sodium borate,
which was necessary to adjust the pH of propolis extract.
Various stocks of propolis from different preparations could be
different in their composition, depending on the plants and flowers
available for bees. Therefore, the antiviral capacity of each stock of
propolis is compared to others by determining the ratio of IC50 (the
concentration required to confer a 50% inhibition of viral infection)
to CC50 (the concentration that causes 50% toxicity to control
uninfected cells).

In Vitro experiments

Viral infection. Monolayers of Vero cells were incubated with

viral suspension (HSV-1 of multiplicity of infection of 1) in RPMI

medium containing 2% newborn calf serum at 378C for 2 hours.
Viral infection was monitored as follows: a) Cytopathic effect: The
unabsorbed virus was removed and fresh medium containing 2%
serum was added to the cultures, which were incubated at 378C
Propolis Activity against Herpes Viruses

923

Original Articles

.
.
.

in humidified air containing 5% CO2 until the end of the

experiment. Each day, the cultures were examined for evidence
of the cytopathic effect, defined as areas of complete destruction
of cells or of morphologically modified cells and expressed as
percentage of damaged cells in the inspected fields. b) Plaque
assay: The unabsorbed virus particles were removed, and fresh
medium containing 0.6% agar and 2% newborn calf serum was
layered over each Vero-cell monolayer. The monolayers were
incubated at 378C in humidified air containing 5% CO2 for 6
days. At the end of the experiment, the overlay was removed, the
cell monolayer was fixed with 10% formalin in saline, and the
cells were stained with crystal violet. The number of plaques was
counted.
Treatment with propolis: Vero cell cultures infected with 1 moi
of HSV-1 were treated with 0.1, 1, 5, 10 or 20% of propolis extract
2 hours before infection, at the time of infection, or 2 hours after
infection. The treatment with propolis extract was terminated 24
hours post-infection. The cultures were maintained at 378C for
10 days.
Treatment with caffeic acid phenethyl ester: Vero cells
infected with 1 moi of HSV-1 were treated with various
concentrations of CAPE 2 hours before infection. The treatment
with CAPE was terminated 24 hours post-infection. The cultures
were maintained at 378C for 10 days.
Toxicity of propolis or CAPE: The toxicity of propolis or CAPE to
cell cultures was examined as follows: Vero cells, seeded at a
concentration of 0.9 x 106 cells per 9.6 cm2 plate, were incubated
at 378C for 24 hours in the absence or presence of various
concentrations of propolis extract or CAPE. The medium was
then replaced with fresh medium and the incubation was
continued for a further 7 days. Each day the cells were examined
for morphologic changes, and the number of cells was counted
with a hemacytometer. The maximum tolerated dose was taken
as the maximum non-toxic concentration.

Animal experiments

Rats: Infection with HSV-1 and treatment with propolis were

performed as follows. Rats were infected with HSV-1
either by scratching small areas of the skin with the
needle of a syringe containing 0.1 ml of HSV-1
suspension or by i.p. inoculation with 0.1 ml of HSV1 suspension. Immediately after inoculation, treatment was started with 0.5 ml of propolis extract, 5%
or 20%, s.c. or i.p. three times a day for 2 weeks.
After completion of treatment, animals were followed for 4 weeks. The number of days to the
appearance of the first symptoms of the disease
(skin vesicles) and of symptoms of encephalitis,
changes in weight or behavior, and time of death
were recorded. Control animals were injected with
PBS (pH 7.2) instead of propolis extract.

m.o.i. = multiplicity of infection

CAPE = caffeic acid phenethyl ester
PBS = phosphate-buffered saline

924

M. Huleihel and V. Isanu

Table 1.

Group of
animals

Rabbits: Rabbits were infected by scratching6 the cornea of one

eye with 0.1 ml of viral suspension (3x10 pfu/ml); the other

(control) eye was scratched with 0.1 ml of PBS. Propolis
treatment was applied locally, as 0.4 ml of a 5 or 20% propolis
solution to both eyes three times a day for up to 2 weeks, or
given i.v. as 0.8 ml of a 5 or 20% propolis solution twice a day for
10 days. The control drug for the ophthalmic experiments was
Zovirax (GlaxoWellcome) applied to the corneas as ointment.
Inflammatory effects, weight changes, alterations in behavior
(agitation and walking around), illness and time of death were
followed for 4 weeks after infection.
Toxicity of propolis to animals: Rats were injected s.c. or i.p.
with 1 ml of 1, 5 or 20% propolis extract twice a day for 2 weeks.
Behavioral alterations, symptoms of illness and changes in
weight were followed for 4 weeks after treatment. For the rabbits,
some were injected i.v. with various concentrations of propolis
once a day for 2 weeks. For the eye tests, the eyes of the rabbits
were scratched, and various concentrations (5 or 20%) of
propolis, as 0.4 ml of drops, were applied three times a day
for 2 weeks. Inflammatory effects, illness, and weight changes
were recorded for 4 weeks post-treatment. Control animals (rats
or rabbits) were injected with PBS.

Statistical analysis

The differences between the various groups of animals that were

infected with or without treatment with propolis extract were
statistically assessed by t-test.
Results

Toxicity of propolis extract

. Cell culture: Most concentrations of propolis extract did not
cause any morphologic modifications, except for the high
concentration of 20%. Similar results were obtained for cell
growth (data not shown).
. Animals: In rats, there was no significant influence of the
treatment with propolis extract on the weight of animals. The
animals did not show any negative side effects, such as illness or

Effect of 5% propolis extract on the development of HSV-1 infection in rats

Non-infected,
non-PE treated
Non-infected,
PE treated
Infected,
non-PE treated
Infected,
PE treated

Appearance
of symptoms
Treatment Days %
route
(p.i.)

Death
Days
(p.i.)

Infective
virus
%

NT

s.c.
i.p.

0
0
98+2

7-9

0
0
98+2

NT
NT
98+2

s.c.
i.p.

56
5

10+0.5
5+0.3

14
10

10+0.5
5+0.3

10+0.5
5+0.3

The number of animals presenting symptoms and subsequently dying was expressed as the percentage
of the total number of tested animals. A total of 40 animals were used for each treatment. Infective virus
(collected in swabs from infected areas 3 days p.i.) was tested in cell culture. The number of animals
containing infective virus was expressed as a percentage of the total number of animals tested.
p.i.= post infection, NT = Not tested. Data are mean + SD (n = 40).

IMAJ . Vol 4, Supplement . November 2002

Original Articles
Table 2.

Effect of 5% propolis extract on the development of HSV-1 infection in rabbit eyes

Group of
animals

Appearance
of symptoms
Treatment Days
%
route
(p.i.)

Death
Days
(p.i.)

Infective
virus
%

Non-infected,

0
NT
non-PE treated
Non-infected,
Drops

0
NT
PE or Zovirax i.v.

0
NT
treated
Infected,
45
100
1012
100
100
non-PE treated
Infected,
Drops
9*
35+2

0
35+2
PE treated
i.v.
10**
30+2

0
30+2
Infected, treated Ointment 78
40+3

0
40+3
with Zovirax
The number of animals containing infected virus was expressed as the percentage of the total
number of animals tested. A total of 10 rabbits were used for each treatment. Infective virus
(collected in swabs from infected eyes 3 days p.i.) was tested in cell culture.
* Light keratitis with secretions
** Reversible light keratitis
Data are mean + SD (n = 10).

inflammation [Table 1; non-infected, PE treated] or behavioral

changes. In rabbits, the animals whose eyes had been scratched
and then treated with propolis extract, in the form of drops, did
not show any inflammatory effects or behavioral modifications
over the whole experiment period [Table 2; non-infected, PE
treated]. Similar results were obtained for rabbits injected i.v.
with propolis extract vs. control animals.

effect of the propolis extract on HSV-1 infection was

evaluated by the plaque assay. The results showed
clearly that there was no direct interaction between
the virus and the propolis extract [Figure 2]. The
approximate 12% reduction in plaque-forming units
associated with 10% propolis extract [Figure 2] is due
to the remaining extract (1% after a dilution of 102) in
the medium during the infection time.
Propolis extract/cell interaction: In order to test for a
possible interaction between propolis and host cells,
which may interfere with virus adsorption, Vero cells
were incubated with a medium containing different
concentrations of propolis extract at 378C for 2 hours.
Unbound propolis was then removed, and cell
monolayers were washed three times with PBS. These
cell cultures were infected with 1 moi of HSV-1 and
the number of plaques formed was determined.
Strong inhibition of viral infection was obtained for
the pretreated cell cultures [Figure 2]. The degree of
inhibition was similar to that obtained when the same
concentration of propolis extract was added at the
time of infection.

Effect of CAPE on HSV-1 infection in cell culture

CAPE, an active component of propolis, was tested for its possible

inhibitory activity against HSV-1 infection of Vero cells in culture.
Cells were treated with various concentrations of CAPE 2 hours
before infection with HSV-1. The treatment was terminated 24 hours
after infection. Our results showed that 10 M of CAPE was able to
inhibit about 70% of plaque formation. Higher concentrations of

Effect of propolis extract on HSV-1 and 2 infection in cell culture

When Vero cells infected with 1 moi of HSV-1 were exposed to 5% of

propolis extract 2 hours before or at the time of HSV-1 inoculation,
the propolis extract conferred almost a complete protection against
the cytopathic effect of HSV-1 [Figure 1]. A minimal protective effect
of 512% was obtained with 0.1% of propolis extract, while the 1%
concentration gave 8590% protection, and 0.5% gave 50%
protection. The addition of 10% propolis extract 2 hours postinfection gave as much as 8085% protection [Figure 1]. Similar
results were obtained when propolis extracts were used against
HSV-2 infection (data not shown). Similar results were obtained
also when Rumanian propolis extract was used as a control in these
experiments (data not shown).
virus absorption by propolis extract
It is possible that propolis prevents HSV-1 infection by blocking its
adsorption into the host cells. To investigate the mode of blocking,
possible interaction of propolis with virus particles and host cells
was tested.
. Propolis/virus interaction: 5 L of 4 x 108 pfu/ml of HSV-1 were
mixed and incubated with 5 L of various concentrations of
propolis extract at 378C for 30 minutes. Thereafter, the effect of
free propolis was reduced by making a 102 dilution of the
mixture with medium containing 2% serum before infection. The
Inhibition of

IMAJ . Vol 4, Supplement . November 2002

Effect of propolis extract on HSV-1 infection in cell

culture. Monolayers of Vero cells were treated with various
concentrations of propolis extract (0.1%, 1%, 5%, 10%) 2 hours
before the infection (&), at the time of infection (*) or 2 hours
post-infection (~) with HSV-1 (1 moi). The percentage of damaged
cells % cytopathic effect (CPE) as compared with the control
(infected but untreated cultures) is plotted vs. the concentration of
propolis extract. Data are mean + SD (n = 4).
Figure 1.

Propolis Activity against Herpes Viruses

925

Original Articles

Effect of propolis extract on infection with HSV-1 of

vero cells. (&) Vero cells were infected with 1 moi of HSV-1 in the
presence of various concentrations of propolis extract. (*) Cells
infected with HSV-1 (1 moi) that had been pretreated with
propolis extract; (~) cells pretreated with propolis extract before
being infected with 1 moi of HSV-1. Plaque forming units (PFU)
were evaluated by the standard plaque assay. Data are mean +
SD (n = 4).
Figure 2.

infected animals that had been treated with 5% or 20% propolis

extract did not show any symptoms of the disease. The
remaining animals showed moderate reversible symptoms 9
10 or 1314 days post-infection (for the 5% and 20% concentrations of propolis extract, respectively). All the animals recovered
completely by 710 days after the appearance of the symptoms
when treatment with propolis extract was continued up to
complete recovery. None of the treated animals died. In the
ophthalmic experiments, protection provided by Zovirax was not
superior to that conferred by the 5% propolis extract [Table 2].
Infective virus was detected in all infected areas in non-treated
animals, but not in treated animals except in those that showed
viral symptoms [Table 2]. Additionally, when 5% of propolis
extract was given locally as drops to rabbit eyes immediately
after the appearance of external symptoms (3 days postinfection), there was an impressive recovery in 80% of the
infected animals (data not shown). The activity of propolis
extract against the development of HSV-1 infection in eyes of
rabbits was statistically significant when compared to the
infected untreated controls, as assessed by t-test (P < 0.001).
Discussion

Our data show a significant anti-HSV-1 effect of the aqueous

propolis extract, which provided very good protection against HSV-1
and 2 (not shown) infection both in vitro and in vivo, namely, in
newborn rats and in rabbits (corneal keratitis) (P < 0.001 as
assessed by t-test).
The most efficient concentration in vivo was 5%. This concentraCAPE were highly toxic to the cells since they inhibited both RNA
tion gave protection without any concomitant toxic or other
and protein synthesis and cell proliferation (data not shown).
negative side effects. In the animal studies, a concentration of
20% was completely effective in preventing the development of
. Rats: When 23 day old rats were treated s.c. or i.p. with 5% of HSV-1 infection in both rats and rabbits. Even at this high
propolis extract immediately post-infection with HSV-1, symp- concentration, there were no toxic or adverse side effects. In the
toms of illness were reduced in almost 90% of the animals, and rabbits, even the lower (5%) concentration completely prevented
death was prevented in almost 95% [Table 1]. In the cases where viral encephalitis and death [Table 1]. The main difference between
propolis did not provide protection, there was, however, a the 5% and 20% treatments was expressed in the longer delay
significant delay in the appearance of symptoms (23 days) and period in the appearance of symptoms for the 20% treatment;
death (26 days), irrespective of the route of propolis adminis- otherwise, the protective effect significant reduction in the
tration [Table 1]. In the animals treated with the 20% propolis intensity of symptoms, with subsequent healing was similar for
extract, the appearance of symptoms and death were almost the two treatments.
The fact that the propolis extract conferred efficient protection
completely prevented (98%) (data not shown). Infective virus was
detected in all infected areas in non-treated animals but not in against HSV-1 infection in vivo, irrespective of the route of
treated animals except in those that showed viral symptoms administration (s.c., i.p., i.v., or topical), points to the excellent
[Table 1]. The activity of propolis extract against the develop- potential of this product as a therapeutic agent for all kinds of
ment of HSV-1 infection in rats was statistically significant when herpetic infections. The lack of toxicity is an additional quality that
compared to the infected untreated controls, as assessed by t- indicates that this product bears the potential to become an antiHSV-1 drug. This conclusion is in keeping with the results of
test (P< 0.001).
. Rabbits: In rabbits whose eyes had been infected with HSV-1, Rumanian and Canadian studies [4,19] in which raw propolis
propolis extract 5% or 20%, given i.v. or locally as drops, proved highly effective in the treatment of HSV-1 and HSV-2
immediately (1520 min) post-infection had a highly efficient infections in humans.
The exact mechanism of antiviral activity of the propolis extract
protective effect [Table 2]; data of 20% propolis treatment is not
shown. All control animals infected with HSV-1 exhibited is still unclear. Our study showed that the addition of propolis
external symptoms (keratitis and secretions) by 34 days after extract to a cell culture 2 hours before or at the time of viral
infection and died as a result of encephalitis 1012 days after infection completely blocked the development of the viral
infection [Table 2]. Most (70% and 90%, respectively) of the infection. This effect could be due to blocking by propolis of
Effect of propolis extract on HSV-1 infection in vivo

926

M. Huleihel and V. Isanu

IMAJ . Vol 4, Supplement . November 2002

Original Articles

the cell membrane receptors for HSV. Interaction of propolis with

the cell membrane could block penetration of viral particles into
the cells and/or could induce internal changes inside the host
cells, which would in turn affect the virus replication cycle. Our
data provide support for this latter possibility of interference with
a late step during the virus replication cycle; we found that the
addition of propolis as long as 2 hours post-infection significantly prevented (~80% inhibition) the development of viral
infection [Figure 1]. Our experiments also showed, however, that
the propolis extract had no direct effect on the virus particles
[Figure 2]. Our results [Table 2] show that activity of propolis
extract against HSV-1 infection could be attributed partially to
CAPE, one of the components of propolis.
It has previously been suggested that the flavonoids present in
propolis interfere with the intracellular redox processes stimulated
by the viral multiplication [1]. Stimulation of the redox processes, in
turn, promotes the production of various free radicals, which have a
noxious influence on cell viability. The function of flavonoids as
scavengers of free radicals [2022] could attenuate the oxidative
stress determined, especially, by oxygen free radicals [21,23,24]
and, consequently, induce an antiviral protection. The exact
mechanism of action of this antiviral activity of the propolis extract
is currently being tested and identified in our laboratory.
Acknowledgment. We thank Ms. I. Mureinik for editorial review of the
manuscript.

8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.

References

1. Amoros M, Simoes CMO, Girre L, Sauvager F, Cormier M. Synergistic

effect of flavons and flavonols against Herpes simplex virus-type 1 in cell
culture. Comparison with the antiviral activity of propolis. J Nat Prod
1992;55:173240.
2. Dumitrescu M, Esanu V, Crisan I. Mecanismes de l'action antiherpetique
de l'extrait aqueux de propolis. I. Action antioxydante sur cultures
fibroblastes humains. Rev Roum Med Virol 1992;43:16573.
3. Negre-Salvayre A, Mabile L, Delchambre Y, Salvayre R. a-tocopherol,
ascorbic acid and rutin inhibit synergistically the copper-promoted LDL
oxidation and the cytotoxicity of oxidized LDL to cultured endothelial
cells. Biol Trace Elem Res 1995;47:8191.
4. Vynograd N, Vynograd N, Sosnowski Z. A comparative multi-centre study
of the efficacy of propolis, acyclovir and placebo in the treatment of
genital herpes (HSV). Photomedicine 2000;7:16.
5. Valcic S, Montenegro G, Mujica AM, et al. Photochemical, morphological, and biological investigations of propolis from central Chile. J Biosci
1999;54:40616.
6. Choi Y, Lee W, Nam S, Choi K, Park Y. Apoptosis induced by propolis
in human hepatocellular carcinoma cell line. Int J Mol Med 1999;4:
2932.
7. Kimoto T, Arai S, Kohguchi M, et al. Apoptosis and suppression of tumor

21.
22.
23.
24.

growth by artepillin c extracted from Brazilian propolis. Cancer Detect

Prev 1998;22:50615.
Vechet L. Propolis effects on some species of microorganisms and fungi.
In: Propolis. Bucharest: Apimondia, 1973.
Volpert R, Elstner E. Biochemical activities of propolis extracts. II.
Photodynamic activities. Z Naturforsch 1993;48c:85862.
Ozturk F, Kurt E, Cerci M, et al. The effect of propolis extract in
experimental chemical corneal injury. Ophthalmic Res 2000;32:1318.
Banskota AH, Tezuka Y, Midorikawa K, Matsushiga K, Kadota S. Two
novel cytotoxic benzofuran derivatives from Brazilian propolis. J Nat Prod
2000;63:12779.
Claus R, Kinscherf R, Gehrke C, et al. Antiapoptotic effects of propolis
extract and propol on human macrophages exposed to minimally
modified low density lipoprotein. Arzneim Forsch 2000;50:3739.
Amoros M, Lourton E, Boustie, Y Girre, L, Sauvager F, Cormier M.
Comparison of the anti-Herpes simplex virus activities of propolis and 3methyl-but-2-enyl caffeate. J Nat Prod 1994;57:6447.
Bojnansky V, Kosljarova V. The inhibitory effect of propolis on some
plant viruses. In: Propolis. Bucharest: Apimondia, 1973.
Cajal N, Esanu V, Crisan I, Ciobanu E. Nivcrisol A Propolis Derived
Product with Antiviral Properties. Bucharest: Institute of Drugs and
Pharmaceutical Research, 1986.
Crisan I, Mutiu A, Sahnazarov N, Cioca V, Esanu V. The propolis effect on
herpes viruses in vitro. Third International Symposium on Apitherapy,
Bucharest, 1976.
Harish Z, Rubinstein A, Golondner M, Elmaliah M, Mizrahi Y.
Suppression of HIV-1 replication by propolis and its immunoregulatory
effect. Drugs Exp Clin Res 1997;23:8996.
Crisan I, Esanu V Patent No. 101730/17.VII. 1980 (Rumania).
Giurcaneanu D, Serbanescu F, Crisan I, Esanu V. Treatment of ocular,
cutaneous and mucosal HSV infection with a propolis aqueous extract.
Third Symposium on Rumanian Drugs, Bucharest, 1985.
Mathiesen L, Wang S, Halvorsen B, Malterud KE, Sund RB. Inhibition of
lipid peroxidation in low-density lipoprotein by the flavonoid myrigalone B and ascorbic acid. Biochem Pharmacol 1996;51:171925.
Robak Y, Grywlewski RJ. Flavonoids are scavengers of superoxide anions.
Biochem Pharmacol 1988;37:83741.
Whalley DF, Hoult CV, Jessup JRS, Leake WDS. Flavonoids inhibit the
oxidative modification of low-density lipoproteins by macrophages.
Biochem Pharmacol 1990;39:14735.
Nakayama T, Yamada M, Osawa T, Kawakishi S. Suppression of active
oxygen-induced cytotoxicity by flavonoids. Biochem Pharmacol 1993;
45:2657.
Tsuda T, Shiga K, Oshima K, Kawakishi S, Osawa T. Inhibition of lipid
peroxidation and the active oxygen radical scavenging effect of
anthocyanin pigments isolated from Ph. vulgaris L. Biochem Pharmacol
1996;52:103339.

Correspondence: Dr. M. Huleihel, Institute for Applied Biosciences,

Ben-Gurion University of the Negev, P.O. Box 653, Beer Sheva 84105,
Israel.
Phone: (972-8) 646-1999
Fax: (972-8) 647-2970
email: mahmoudh@bgumail.bgu.ac.il

What's important is what we become, not what we have. It's how well we've
learned to serve. It's the relationships we've built.

Anonymous

IMAJ . Vol 4, Supplement . November 2002

Propolis Activity against Herpes Viruses

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