Induction of p21 protein protects against
sulforaphane-induced mitotic arrest in
LNCaP human prostate cancer cell line
Anna Herman-Antosiewicz,
1
Hui Xiao,
2
Karen L. Lew,
2
and Shivendra V. Singh
2
1
Department of Molecular Biology, University of Gdan ´sk,Gdan ´sk,
Poland and
2
Department of Pharmacology and University of
Pittsburgh Cancer Institute, University of Pittsburgh School of
Medicine, Pittsburgh, Pennsylvania
Abstract
Previous studies have indicated that D,L-sulforaphane
(SFN), a synthetic cancer chemopreventive analogue of
cruciferous vegetable-derived isomer (À)-1-isothiocya-
nato-(4R )-(methylsulfinyl)-butane, activates a checkpoint
kinase 2 (Chk2)–dependent G
2
-M phase cell cycle arrest
inp53-deficienthumanprostatecancercells.Becausep53
is a downstream target of Chk2 kinase and known to
regulate G
2
-M transition by transcriptional regulation of
cyclin-dependent kinase (Cdk) inhibitor p21
Cip1/Waf1
(p21),
the present study was undertaken to determine the role of
p21inSFN-inducedcellcyclearrestusingwild-typep53–
expressing cell line LNCaP. The SFN treatment caused a
modestincreaseinSphasefractionandamarkedincrease
in G
2
-M fraction in LNCaP cells in a concentration- and
time-dependent manner. The SFN-induced S phase arrest
correlated with a reduction in protein levels of cyclin D1,
cyclin E, Cdk4, and Cdk6, whereas activation of the G
2
-M
checkpoint was accompanied by induction of cyclin B1
and down-regulation of Cdk1 and Cdc25C protein levels.
The SFN-treated LNCaP cells were also arrested in mitosis
as revealed by immunofluorescence microscopy and
increased Ser
10
phosphorylation of histone H3, a sensitive
marker for mitotic cells. The SFN treatment increased
activating phosphorylation of Chk2 (Thr
68
) that was
accompanied by induction of p53 and p21. The SFN-
induced mitotic arrest was statistically significantly
increased by small interfering RNA–based knockdown of
p21. However, p21 protein knockdown did not have any
appreciable effect on SFN-induced cytoplasmic histone-
associated DNA fragmentation (apoptosis). In conclusion,
the present study indicates that induction of p21 protects
against SFN-induced mitotic arrest in LNCaP cells. [Mol
Cancer Ther 2007;6(5):1673–81]
Introduction
Epidemiologic studies continue to support the premise
that dietary intake of cruciferous vegetables may lower the
risk of different types of malignancies including prostate
cancer (1 – 4). Anticarcinogenic effect of cruciferous vege-
tables is attributed to organic isothiocyanates, which are
generated due to myrosinase-mediated hydrolysis of
corresponding glucosinolates (5, 6). Broccoli is a rather
rich source of the isothiocyanate compound (À)-1-isothio-
cyanato-(4R )-(methylsulfinyl)-butane (L-SFN). The L-SFN
and its synthetic analogue D,L-sulforaphane (SFN) have
generated a great deal of research interest due to their
remarkable anticancer effects (7 – 14). For instance, L-SFN
or SFN has been shown to afford significant protec-
tion against chemically induced cancer in animal models
(8, 11 – 13). Cancer chemoprevention by L-SFN or SFN has
been observed against 9,10-dimethyl-1,2-benzanthracene –
induced mammary cancer in rats, azoxymethane-induced
colonic aberrant crypt foci in rats, benzo[a ]pyrene-induced
forestomach cancer in mice, and 7,12-dimethylbenz(a )an-
thracene/12-O -tetradecanoylphorbol 13-acetate – induced
skin tumorigenesis in mice (8, 11 – 13). More recently,
SFN administration was shown to significantly inhibit
lung metastasis induced by B16F-10 melanoma cells in
C57BL/6 mice (14).
Evidence is accumulating to indicate that SFN can
suppress proliferation of cultured cancer cells by causing
cell cycle arrest and apoptosis induction (15 – 24). Interest-
ingly, normal epithelial cells, including a normal human
prostate epithelial cell line PrEC and an immortalized
normal human bronchial epithelial cell line BEAS-2B, are
significantly more resistant to apoptosis induction by SFN
compared with cancer cells (21). Consistent with these
cellular results, oral administration of SFN significantly
retards growth of PC-3 human prostate cancer xenografts
in nude mice without causing weight loss or any other side
effects (18). Inhibition of histone deacetylase activity by
SFN has also been observed in human benign prostate
hyperplasia epithelial cell line BPH-1 and PC-3 and LNCaP
human prostate cancer cells (23).
An understanding of the mechanism(s) by which SFN
causes cell cycle arrest and apoptosis induction is critical
for its further clinical development because this knowledge
Received 12/28/06; revised 3/26/07; accepted 3/30/07.
Grant support: National Cancer Institute/USPHS grants CA115498 and
CA101753.
The costs of publication of this article were defrayed in part by the
payment of page charges. This article must therefore be hereby marked
advertisement in accordance with 18 U.S.C. Section 1734 solely to
indicate this fact.
Note: A. Herman-Antosiewicz and H. Xiao contributed equally to this
work.
Requests for reprints: Shivendra V. Singh, 2.32A Hillman Cancer Center
Research Pavilion, University of Pittsburgh Cancer Institute,
5117 Centre Avenue, Pittsburgh, PA 15213. Phone: 412-623-3263;
Fax: 412-623-7828. E-mail: singhs@upmc.edu
Copyright C 2007 American Association for Cancer Research.
doi:10.1158/1535-7163.MCT-06-0807
1673
Mol Cancer Ther 2007;6(5). May 2007
Research.
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