The Novel Triterpenoid C-28 Methyl Ester of 2-Cyano-3,
12-Dioxoolen-1, 9-Dien-28-Oic Acid Inhibits Metastatic Murine
Breast Tumor Growth through Inactivation of STAT3 Signaling
Xiaoyang Ling,
1
Marina Konopleva,
1
Zhihong Zeng,
1
Vivian Ruvolo,
1
L. Clifton Stephens,
2
Wendy Schober,
1
Teresa McQueen,
1
Martin Dietrich,
1
Timothy L. Madden,
3
and Michael Andreeff
1
1
Section of Molecular Hematology and Therapy, Department of Stem Cell Transplantation and Cellular Therapy,
2
Department of Veterinary
Medicine and Surgery, and
3
Department of Experimental Therapeutics, The University of Texas M. D. Anderson Cancer Center,
Houston, Texas
Abstract
We and others have reported that C-28 methyl ester of 2-
cyano-3, 12-dioxoolen-1, 9-dien-28-oic acid (CDDO-Me) effec-
tively inhibits the growth of multiple cancer cell types. Our
previous studies indicated that prolonged CDDO-Me treat-
ment inactivated extracellular signal-regulated kinase signal-
ing in acute myelogenous leukemia cells. Whether treatment
with CDDO-Me has an earlier effect on other proteins that are
important for either signal transduction or oncogenesis is
unknown. Constitutively activated signal transducer and
activator of transcription 3 (STAT3) is frequently found in
human breast cancer samples. Constitutively activated STAT3
was shown to up-regulate c-Myc in several types of cancer and
has a feedback effect on Src and Akt. To examine the effects of
CDDO-Me on STAT3 signaling in breast cancer, we used the
murine 4T1 breast tumor model, which is largely resistant to
chemotherapy. In vitro , after treatment of 4T1 cells with 500
nmol/L CDDO-Me for 2 h, we found (a ) inactivation of STAT3,
(b) inactivation of Src and Akt, (c ) 4-fold reduction of c-Myc
mRNA levels, (d ) accumulation of cells in G
2
-M cell cycle
phase, (e ) abrogation of invasive growth of 4T1 cells, and ( f )
lack of apoptosis induction. In in vivo studies, CDDO-Me
completely eliminated 4T1 breast cancer growth and lung
metastases induced by 4T1 cells in mice when treatment
started 1 day after tumor implantation and significantly
inhibited tumor growth when started after 5 days. In vivo
studies also indicated that splenic mature dendritic cells were
restored after CDDO-Me treatment. In summary, these data
suggest that CDDO-Me may have therapeutic potential in
breast cancer therapy, in part, through inactivation of STAT3.
[Cancer Res 2007;67(9):4210–8]
Introduction
Signal transducer and activator of transcription 3 (STAT3) has
frequently been found in human breast cancer samples (1). STAT3-
mediated up-regulation of c-Myc levels was proposed to be one of
the important mediators of STAT3 oncogenesis (2). c-Myc is a
potent oncoprotein that has been found to play a critical role
in tumorigenesis (3). The use of an inducible c-Myc model
showed tumor growth or suppression, respectively, by controlling
c-Myc expression in a murine hepatocarcinoma model (4). Our
previous studies and that of others have shown that c-Myc
expression is regulated by constitutive activation of STAT3 in
mouse breast cancer and other types of cancer cells (5–7). As
the immediate upstream regulator of STAT3, Src has been
considered to play an important role in breast carcinogenesis.
However, we found that Src can be inactivated by STAT3
knockdown in 4T1 cells (5). We have also shown that Akt can
be inactivated following STAT3 knockdown in 4T1 cells (5),
suggesting that STAT3 is required for maintaining the activation
status of Src and Akt. As a downstream target of Src, Akt plays
an important role in regulating antiapoptotic proteins and
proteins that regulate cell cycle progression (8). However, it is
unclear how these intracellular signal transduction proteins
interact with each other, and the biological consequences of
these interactions remain uncertain. Recent reports provided
convincing evidence that constitutively activated STAT3 in breast
cancer cells is associated with impaired cell-mediated immunity
(9) and that STAT3 activation in cancer cells may allow them to
evade host immune surveillance (10). In agreement with these
observations, we reported that knockdown of STAT3 completely
abolished breast tumor formation in an immunocompetent
mouse model (5).
The 4T1 cell line was originally derived from a spontaneous
mammary carcinoma in BALB/c mice (11). It has been reported
that 4T1 cells mimic the effects of human mammary carcinoma
in that morbidity is due to the outgrowth of spontaneous
micrometastatic tumor cells that migrate to distant organs
relatively early during primary tumor growth (12, 13) and are
notoriously chemoresistant (14). Therefore, 4T1-driven breast
cancer in BALB/c mice constitutes an appropriate in vivo system
for modeling human breast cancer with regard to tumor growth
and metastasis. As such, the 4T1/BALB/c mouse system also
provides a useful experimental model for exploring the biological
effects in mammary tumorigenesis and for studying the effects
of small molecules that target signal transduction pathways
in vivo .
2-Cyano-3, 12-dioxoolen-1, 9-dien-28-oic acid (CDDO) and its
derivative CDDO-Me are synthetic triterpenoids proven to be
effective agents in controlling cancer cell growth in preclinical
models of leukemias, multiple myeloma, lymphomas, and solid
tumors, such as breast, pancreatic, and colon cancer (15–18).
CDDO-Me has been shown by us to (a ) promote apoptosis in
several leukemic cell lines, (b) induce proapoptotic Bax protein as a
Requests for reprints: Michael Andreeff, Department of Blood and Marrow
Transplantation, Unit #448, 1515 Holcombe Boulevard, The University of Texas M. D.
Anderson Cancer Center, Houston, TX 77030-4009. Phone: 713-792-7260; Fax: 713-794-
4747; E-mail: mandreef@mdanderson.org.
I2007 American Association for Cancer Research.
doi:10.1158/0008-5472.CAN-06-3629
Cancer Res 2007; 67: (9). May 1, 2007 4210 www.aacrjournals.org
Research Article
Research.
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