Mini-Reviews in Organic Chemistry, 2008, 5, 171-178 171
1570-193X/08 $55.00+.00 © 2008 Bentham Science Publishers Ltd.
Antagonists Against Anti-Apoptotic Bcl-2 Family Proteins for Cancer
Treatment
Jignesh M. Doshi and Chengguo Xing*
Department of Medicinal Chemistry, College of Pharmacy, University of Minnesota, 8-101 WDH, 308 Harvard St S. E.,
Minneapolis MN 55455, USA
Abstract: Apoptosis is a programmed cell death process, critical for normal cellular development and tissue homeostasis.
B-cell lymphocyte/leukemia 2 (Bcl-2) family proteins are important regulators of apoptosis. Numerous studies have dem-
onstrated that over-expressing anti-apoptotic Bcl-2 proteins is one mechanism for cancer cells to acquire resistance against
cancer chemotherapies, suggesting antagonizing these proteins would be a potential approach to overcoming such drug re-
sistance. This review briefly discusses the principle and the recent advances in the development of such antagonists, with
some highlights about several promising antagonists.
Keywords: Bcl-2 antagonists, drug resistance, apoptosis, cancer.
INTRODUCTION
Apoptosis is a highly regulated and energy-dependent
process of cellular suicide that can be triggered by a multi-
tude of internal and external stimuli [1]. Apoptosis plays an
important role in many developmental processes as well as
maintenance of tissue homeostasis. Indeed, impaired apopto-
sis is a hallmark of various cancers [2]. As many cancer
therapies eliminate tumor cells through induction of apopto-
sis directly or indirectly, impairment in apoptosis also leads
to drug resistance to cancer therapies [3].
REGULATION OF LIFE AND DEATH BY THE BCL-2
FAMILY PROTEINS
One mechanism for cancers to acquire resistance to apop-
tosis is through the over-expression of anti-apoptotic Bcl-2
family proteins, a subfamily of Bcl-2 family proteins. Bcl-2
family proteins comprise of over 20 members, which are the
crucial regulators of apoptosis [4]. The Bcl-2 family proteins
can be functionally classified into two broad subfamilies.
The first subfamily comprise of family members that protect
a cell from apoptosis, named as anti-apoptotic Bcl-2 proteins
(Bcl-2, Bcl-X
L
, Bcl-w, Mcl-1, A-1, and Bcl-B) [5, 6]. The
*Address correspondence to this author at the Department of Medicinal
Chemistry, College of Pharmacy, University of Minnesota, 8-101 WDH,
308 Harvard St S. E., Minneapolis MN 55455, USA; Tel: 612-626-5675;
Fax: 612-624-0139; E-mail: xingx009@umn.edu
other subfamily consists of family members that promote
apoptosis, named pro-apoptotic Bcl-2 proteins. These two
subfamilies antagonize each other and the balance between
them determines the cell’s fate to undergo apoptosis or to
survive. Most of the anti-apoptotic members contain four
conserved motifs named as Bcl-2 homology (BH) domains
(BH1 – BH4) except Mcl-1 contains only three conserved
domains (BH1 – BH3) and lacks the BH4 domain [7] (Fig.
1). The pro-apoptotic Bcl-2 proteins can be further classified
into two subsets. Some members (Bax, Bak, and Bok) have
three BH domains (BH1, BH2, and BH3) while other mem-
bers only have BH3 domain, named as BH3-domain only
pro-apoptotic Bcl-2 proteins (Bik, Bid, Bad, Bim, Bmf, Hrk,
Noxa, and Puma) [8]. Structural studies have revealed that
the BH1, BH2, and BH3 domains in the anti-apoptotic pro-
teins are folded to form an elongated hydrophobic cleft. This
cleft serves as the binding site to interact with the pro-
apoptotic members through their BH3 domain, forming het-
erodimer [9, 10]. Heterodimerization of a pro-apoptotic Bcl-
2 family member and an anti-apoptotic Bcl-2 family member
is one proposed mechanism for these two subfamilies to an-
tagonize each other. The anti-apoptotic Bcl-2 family proteins
engage pro-apoptotic Bcl-2 proteins and keep them in check,
thereby preventing apoptosis. The pro-apoptotic Bcl-2 fam-
ily proteins, when in excess or activated (not bound by anti-
apoptotic ones), can permeabilize the mitochondrial mem-
brane, resulting in the leakage of cytochrome c and other
molecules involved in caspase activation, which leads to
apoptosis [11, 12].
Fig. (1). Schematic representation of structural domains of the Bcl-2 family proteins.