0012-4966/05/0910- © 2005 Pleiades Publishing, Inc. 0410
Doklady Biological Sciences, Vol. 404, 2005, pp. 410–412. Translated from Doklady Akademii Nauk, Vol. 404, No. 5, 2005, pp. 704–706.
Original Russian Text Copyright © 2005 by Solov’eva, Solov’ev, Faskhutdinova, Kudryavtsev, Akatov, Chailakhyan.
The antioxidant properties of thiol-containing com-
pounds are well known [1]. It is established that antiox-
idants combined with metals of variable valence may
form prooxidant catalytic systems. An example of such
a system is ascorbate combined with organic com-
plexes of copper or cobalt, such as vitamin B
12b
or
cobalt phthalocyanines, which were used as antitumor
agents during the past years [2–7]. However, it is
unclear how thiols may serve as prooxidants in combi-
nation with vitamin B
12b
. Antioxidant therapy based on
vitamin–mineral complexes is widely used; hence, the
study of the prooxidant effect of the combination of thi-
ols and vitamin B
12b
is of considerable importance.
In this work, we show that thiol antioxidants com-
bined with vitamin B
12b
form a prooxidant catalytic
system that generates hydrogen peroxide and synergis-
tically induces oxidative stress and apoptotic cell death
of tumor cells.
Epidermoid carcinoma of larynx (HEp-2) cells
obtained from the All-Russian Collection of Cell Cul-
tures (Institute of Cytology, Russian Academy of Sci-
ences, Saint-Petersburg) were grown on DME (Sigma)
containing 10% fetal calf serum (HyClone), 80 μ g/ml
gentamicin, 2.2 g/l sodium bicarbonate and maintained
at 37°C and 5% CO
2
. The substances studied were
added 24 h after plating 5000 cells/well in a 96-well
microplate. The chelators of iron ions deferoxamine
and phenanthroline (Sigma) were added 2 or 0.5 h
before the addition of B
12b
combined with GSH, NAC,
and DTT (MPBiomedicals). Catalase and pyruvate
(Sigma) were added immediately before these sub-
stances. The cells were stained using crystal violet and
studied by means of a photometer to assess the cyto-
toxic effects of these agents [8]. Cytotoxicity was
assessed by the ratio between the amount of living cells
in the experiment and control 48 h after the 2-h incuba-
tion of the cells with the agents studied.
The aberrant distribution of chromatin, which is a
sign of apoptotic cell death, was assessed with the use
of luminescent microscopy using 1 μ g/ml Bisbenzim-
ide H33342 and ethidium bromide (Sigma). Double
staining determines aberrant distributions of chromatin
in living and dead cells (green and red luminescence,
respectively) [9]. The existence of apoptotic cells was
assessed by the presence of particles in sub-G
1
region
on DNA-cytograms obtained by means of flow cytom-
etry using a PARTEC III cytometer [10]. In addition,
apoptosis was assessed by DNA fragmentation with the
use of electrophoresis in 1.5% agarose gel [11].
The concentration of hydrogen peroxide in the
medium was measured using polarographic method by
oxygen concentration after the addition of catalase. The
agents studied were added 24 h after the plating
350 000 HEp-2 cells in T25 flasks. The calibration was
performed by addition of H
2
O
2
into the experimental
chamber after catalase [7].
The oxidative activity of the cells was assessed with
the use of the fluorescent probe 2',7'-dichlorofluores-
cein diacetate (DCHFDA, Molecular probes). The cells
were plated in T25 flasks (350 000 cells per 5 ml of cul-
ture medium). B
12b
and NAC were added a day after
plating. After 2 h of incubation, the cells were detached
by trypsinization, resuspended into fresh culture
medium, and stained with 20 μ M DCFHDA for 30 min.
Then, the cells were washed from the dye and resus-
pended at a concentration of 500 000 cells/ml in cold
PBS. The fluorescence was recorded using a PARTEC
III flow cytometer [12].
We found that the thiols studied (GSH, NAC, and
DTT) had a synergistic cytoxic effect in combination
with vitamin B
12b
. Addition to the cell cultures of the
combinations of thiols and B
12b
for 2 h suppressed cell
proliferation in a dose-dependent manner and initiated
cell death (Fig. 1). The death of cells exposed to the
toxic combinations for 2 h began 6 h later and reached
95% by the end of the first day. Vitamin B
12b
(as much
as 3 mM) or thiols (10 mM or more) added separately
were not toxic for HEp-2 cells after a 2-h incubation. A
similar synergism of the cytotoxic effect of thiols in
combination with vitamin B
12b
was also found during
permanent exposure of the cells to the agents.
The cell death induced by combinations of thiols
with B
12b
was accompanied by signs typical of apopto-
sis [11]. A study of cell death type according to mor-
phologic property showed an increase in the number of
CELL
BIOLOGY
Prooxidant and Cytotoxic Effects of Thiols Combined
with Vitamin B
12b
M. E. Solov’eva, V. V. Solov’ev, A. A. Faskhutdinova, A. A. Kudryavtsev, V. S. Akatov,
and Corresponding Member of the RAS L. M. Chailakhyan
Received June 2, 2005
Institute of Theoretical and Experimental Biophysics,
Russian Academy of Sciences, ul. Institutskaya 3, Pushchino,
Moscow oblast, 142292 Russia