ICANCER RESEARCH 56. 2745-2747. June 15. 19%|
Advances in Brief
Inhibition of Fatty Acid Synthesis Induces Programmed Cell Death in Human
Breast Cancer Cells1
Ellen S. Pizer, Christian Jackisch, Fawn D. Wood, Gary R. Pasternack, Nancy E. Davidson, and Francis P. Kuhajda2
Division of Molecular Pathology. Department of Pathology ¡E.S.P.. F.D.W., G.R.P.. F.P.K.], ami Department of Oncology ¡C.J.,N.E.D.I. The Johns Hopkins Medical Institutions,
Baltimore. Man-land 212X7
Abstract
One of the key limiting factors in the treatment of advanced stage
human epithelial malignancies is the lack of new, selective molecular
targets for antineoplastic therapy. A substantial subset of human breast,
ovarian, endometrial, coloréela!,and prostatic cancers express elevated
levels of fatty acid synthase, the major enzyme required for endogenous
fatty acid biosynthesis, and carcinoma lines are growth inhibited by
cerulenin, a noncompetitive inhibitor of fatty acid synthase. We have
shown previously that the difference in fatty acid biosynthesis between
cancer and normal cells is an exploitable target for metabolic inhibitors in
the in vitro setting and in vivo in a human ovarian carcinoma xenograft in
nude mice. Here, we report that cerulenin treatment of human breast
cancer cells inhibits fatty acid synthesis within 6 h after exposure, that loss
of clonogenic capacity occurs within the same interval, and that DNA
fragmentation and morphological changes characteristic of apoptosis
ensue.
Introduction
The treatment of human epithelial malignancies is limited by drug
resistance and toxic side effects of therapy, which contribute to
ultimate treatment failure for the majority of advanced stage cancer
victims. Identification of new, selective molecular targets for antine
oplastic therapy provides an opportunity for therapeutic advancement.
Although disordered intermediary metabolism in cancer cells has been
known for the better part of this century ( 1), little attention has been
paid to fatty acid metabolism. One early study showed elevated levels
of fatty acid synthesis in tumor tissues, although the significance of
the observations was not appreciated (2). We and others (3-10) have
recently shown that some clinical human ovarian, endometrial, breast,
colorectal, and prostatic cancers overexpress FAS.3 Tumor cells
which express high levels of fatty acid synthesizing enzymes use
endogenously synthesized fatty acids for membrane biosynthesis and
also appear to export large amounts of lipid.4 In marked contrast,
normal cells preferentially utilize dietary lipids.
We have shown recently that these biochemical differences provide
a selective target for metabolic inhibitors both in vitro and in vivo.
Key observations are that: (a) inhibition of the ß-ketoacylsynthase
site of FAS is selectively cytotoxic to cancer cells with increased fatty
Received 2/12/96: accepted 4/25/96.
The cosls of publication of this article were defrayed in part by Ihe payment of page
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' Supported by grants from the National Cancer Institute (CA57545), the American
Institute for Cancer Research, the W. W. Smith Charitable Trust, and the Susan G. Komen
Foundation. E. P. was supported by USPHS Grant T32 AI 07247 and by a grant from Ihe
Steiler Research Fund. C. J. was supported by a fellowship from the Deutsche Forschungs-
gemeinschap.
2 To whom requests for reprints should be addressed, at Department of Pathology. The
Johns Hopkins Medical Institutions, 720 Rutland Avenue. Ross 512, Baltimore, MD
21205.
1The abbreviation used is: FAS, fatty acid synthase.
4 H. S. Heine. F. P. Kuhajda. F. D. Wood, A. Kayler, and G. R. Pasternack, Increased
rates of fatty acid synthesis in breast cancer and identification of unique products,
manuscript in preparation.
acid biosynthesis but not to normal skin fibroblasts in vitro; (b)
addition of palmitate, the major direct product of FAS, reverses the
cytotoxic effects of its inhibition, demonstrating mechanistic spec
ificity (11-14); (c) elevations in fatty acid synthase expression in
carcinoma cell lines are comparable to levels in primary human
tumors assessed by immunohistochemistry; (d) fatty acid synthetic
activities of a carcinoma line grown in vitro or as a murine
xenograft are similar, and are 4- to >20-fold higher than normal
murine tissues; and (e) treatment with the specific FAS inhibitor,
cerulenin, produces regression of established ascites tumor, reduc
tion in ascites incidence, delay in onset of ascites, and significantly
increased survival in a nude mouse xenograft model of human
ovarian carcinoma (15).
The mechanism by which inhibition of fatty acid synthesis pro
duces its antitumor effect remains unexplained. The evidence pre
sented here indicates that exposure of breast carcinoma cells to
cerulenin produces inhibition of fatty acid synthesis within 6 h and
produces irreversable lethal injury measured as a reduction of clono
genic capacity in the same interval. Treated breast carcinoma cells
subsequently undergo DNA fragmentation and morphological
changes characteristic of apoptosis.
Materials and Methods
Cell Lines and Culture Conditions. ZR-75-1, SKBR3, and MCF-7 cells
were maintained in RPMI 1640 with 10% fetal bovine serum except as
otherwise indicated. Cultures were incubated at 37°Cin a humidified 5% CO2
atmosphere. A 5 mg/ml stock solution of cerulenin (Sigma Chemical Co.) in
DMSO was diluted into experimental cultures to final concentrations of 5 or 10
fig/ml.
Measurement of Endogenous Lipid Synthesis. ZR-75-1 cells were
plated at 2 X IO5 cells/well in 24-well plates and incubated overnight prior to
use. Cerulenin (Sigma Chemical Co.) in DMSO was added for indicated
incubation times. Each well was pulse labeled with 0.1 /xCi [U-'4C]acetate
during the final 2 hours of drug treatment and washed, and cellular lipids were
extracted (13) and assayed for I4C by scintillation counting. Controls consisted
of cells incubated with DMSO alone. All determinations were performed in
triplicate with error bars representing the SE.
Clonogenic Assay. After overnight incubation, 1 X 10" ZR-75-1, SKBR3,
or MCF-7 cells were exposed to cerulenin (Sigma Chemical Co.) in DMSO
(0.1-0.2%) for 6 h, washed, detached by trypsin digestion, counted, and
replated at 1000 or 500 cells/60-mm plate in triplicate. Colonies were stained
with crystal violet and counted 3-5 days after plating. Controls consisted of
cells incubated with DMSO without cerulenin. Error bars represent the SE.
DNA Fragmentation Assay and Morphological Evaluation. Exponen
tially growing ZR-75-1 cells were plated at 1-5 x IO4 cells/cm2 in DMEM
with 5% fetal bovine serum and 2 mM glutamine. After attachment, the
medium was changed, and cells were incubated with or without cerulenin
continuously until harvesting. At harvest, medium and trypsinized cells were
combined, and DNA was extracted from pelleted cells and subjected to pulse
field gel electrophoresis as described previously (16). Cells treated in parallel
were spun onto glass slides, fixed in methanol, and stained with Hoechst 33342
for evaluation of morphological changes of apoptosis by fluorescence micros
copy at 480 nm.
2745
Research.
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