(CANCER RESEARCH 58. 717-723. February 15. 1998)
Novel Triterpenoids Suppress Inducible Nitric Oxide Synthase (iNOS) and Inducible
Cyclooxygenäse(COX-2) in Mouse Macrophages1
Nanjoo Suh, Tadashi Honda, Heather J. Finlay, Aaron Barchowsky, Charlotte Williams, Nicole E. Benoit,
Qiao-wen Xie, Carl Nathan, Gordon W. Gribble, and Michael B. Sporn2
Department of Pharmacology and Norris Cotton Cancer Center, Dartmouth Medical School [N. S.. A. B., C. W.. N. E. B., M. B. SJ and Department of Chemistry. Dartmouth
College, Hanover, New Hampshire 03755 ¡T.H., H. J. F., G. Vi.G.], and Department of Médiane, Cornell University Medical College, New York, New York 10021 ¡Q-w.X.,
C.N.J
ABSTRACT
We have synthesized more than 80 novel triterpenoids, all derivatives of
oleanolic and ursolic acid, as potential anti-inflammatory and chemopre-
ventive agents. These triterpenoids have been tested for their ability to
suppress the de novo formation of two enzymes, inducible nitric oxide
synthase (¡NOS) and inducible cyclooxygenase (COX-2), using IFN-y-
stimulated primary mouse macrophages or lipopolysaccharide (1,1'Si-
activated RAW 264.7 macrophages as assay systems. Two synthetic
oleananes, 3,12-dioxoolean-l-en-28-oic acid (TP-69) and 3,11-dioxoolean-
l,12-dien-28-oic acid (TP-72), were highly active inhibitors of de novo
formation of both ¡NOSand COX-2. Both TP-69 and TP-72 blocked the
increase in iNOS or COX-2 mRNA induced by IFN-y or LPS. In addition,
TP-72 suppressed NT-Kit activation in primary macrophages treated with
the combination of II N-y and LPS or II N-y and tumor necrosis factor.
The 3-«(axial)-epimer of ursolic acid suppressed de novo formation of
COX-2, in contrast to naturally occurring 3-ß( equatorial )-ursolic acid.
Inhibitory effects of TP-69 or TP-72 on ¡NOSformation were not blocked
by the glucocorticoid receptor antagonist RU-486, indicating that these
triterpenoids do not act through the glucocorticoid receptor, nor does
TP-72 act as an ¡NOSor COX-2 enzyme inhibitor when added to RAW
cells in which synthesis of these two enzymes in response to LPS has
already been induced. It may be possible to develop triterpenoids as useful
agents for chemoprevention of cancer or other chronic diseases with an
inflammatory component.
INTRODUCTION
One of the major needs in cancer prevention is the development of
effective and safe new agents for chemoprevention. In particular,
there is a unique need for chemopreventive agents targeted at mech
anisms known to be involved in the process of carcinogenesis ( 1). In
recent years, there has been a resurgence of interest in the study of
mechanisms of inflammation that relate to carcinogenesis and in the
use of such mechanisms as the basis for development of new chemo
preventive agents.
The concept that inflammation and carcinogenesis are related phe
nomena has been the subject of many previous studies that have
attempted to link these two processes in a mechanistic fashion (2-4).
The enzymes that mediate the constitutive synthesis of NO and
prostaglandins from arginine and arachidonate, respectively, have
relatively little significance for either inflammation or carcinogenesis.
In contrast, iNOS3 (EC 1.14.13.39) and inducible cyclooxygenase
(COX-2; EC 1.14.99.1) have critical roles in the response of tissues to
Received 10/14/97: accepted 12/19/97.
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.
' This research was supported by Morris Cotton Cancer Center Grant CA-23108. NIH
Grant ROI AI 34543. the Oliver S. and Jennie R. Donaldson Charitable Trust, and
DOD/AMRD Award 17-96-1-6163.
- To whom requests for reprints should be addressed, at Department of Pharmacology
and Toxicology. Dartmouth Medical School. 7650 Remsen, Hanover, NH 03755. Phone:
(603)650-6557: Fax: (603)650-1129; E-mail: Michael.Sporn@danmouth.edu.
1The abbreviations used are: iNOS, inducible nitric oxide synthase; TGF. transform
ing growth factor; COX. cyclooxygenase; OA. oleanolic acid; UA. ursolic acid; PG.
prostaglandin; LPS. lipopolysaccharide; TNF, tumor necrosis factor; NF, nuclear factor.
injury or infectious agents. These inducible enzymes are essential
components of the inflammatory response, the ultimate repair of
injury, and carcinogenesis (5-10). Although physiological activity of
iNOS and COX-2 may provide a definite benefit to the organism,
aberrant or excessive expression of either iNOS or COX-2 has been
implicated in the pathogenesis of many disease processes, as diverse
as septic shock, cardiomyopathy, acute and chronic neurodegenerative
disease, rheumatoid arthritis, and carcinogenesis (11-19).
Immense effort has been devoted to developing new molecules that
are direct inhibitors of the enzymatic activity of either iNOS or
COX-2. However, an alternative approach is to find new agents that
can prevent expression of the respective genes coding for these
enzymes. Glucocorticoids and TGF-ßare such molecules; they both
suppress transcription or translation of iNOS and COX-2 genes (20-
25). A rationale thus exists to develop more selective agents for
suppression of genes that might be overexpressed during the inflam
matory or carcinogenic process. In this report, we attempt to apply this
strategy for the development and evaluation of new triterpenoids.
Although triterpenoids are widely used for medicinal purposes in
many Asian countries, this class of molecules, which resemble ste
roids in their chemical structure, biogenesis, and pleiotropic actions,
has not impacted on the practice of Western medicine. Triterpenoids,
like the steroids, are formed in nature by the cyclization of squalene,
with the retention of all 30 carbon atoms in molecules such as OA and
UA (Fig. 1). Although OA and UA are known to have numerous
pharmacological activities, the potency of these naturally occurring
molecules is relatively weak. Chemical synthesis of new steroid
analogues has provided many useful derivatives that are more potent
and specific than natural parent structures. With this as a model and
considering the known anti-inflammatory and anticarcinogenic activ
ities of OA and UA (26-29), we have synthesized (30) and charac
terized a new series of synthetic triterpenoid analogues as potential
inhibitors of inflammation and carcinogenesis using suppression of
the formation of nitric oxide and prostaglandins as assay systems. We
report here the structures and activities of three of the most potent of
these derivatives of OA and UA. These new agents are significantly
more active than their parent molecules.
MATERIALS AND METHODS
Reagents
Details of the synthesis of TP-69 and TP-72 (see Fig. I for structures) have
been published (30). TP-52 (3-a-OH UA) was synthesized by Jones oxidation
of UA. followed by Meenvein-Ponndorf reduction. Recombinant mouse IFN-y
(LPS content. <10 pg/ml) was purchased from Genzyme (Cambridge, MA);
NF-«B oligonucleotide was purchased from Promega Corp. (Madison, WI);
goat polyclonal COX-1. COX-2 IgG. and anti-goat IgG peroxidase-conjugated
secondary antibody were purchased from Santa Cruz Biotechnology (Santa
Cruz, CA); TGF-ß, was from R&D (Minneapolis. MN); and enzyme immu-
noassay reagents for PGE2 assays were from Cayman Co. (Ann Arbor. MI).
TNF-a was provided by Dr. Jan Vilcek (New York University Medical Center,
New York, NY). LPS (from Escherichia coli 0111:B4 y-irradiated) and all
other chemicals were purchased from Sigma Chemical Co. (St. Louis, MO).
717
Research.
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