Mini Reviews in Medicinal Chemistry, 2001, 1, 5-16 5
Modifying TNFα for Therapeutic Use: A Perspective on the TNF Receptor
System
Akihiro Hasegawa, Watrau Takasaki**, Mark I. Greene and Ramachandran
Murali *
Department of Pathology and Laboratory Medicine, 252 John Morgan Building, University of
Pennsylvania School of Medicine, 36th and Hamilton Walk, Philadelphia, PA 19104, U.S.A.
Abstract: TNFα is an inflammatory mediator that is relevant to several autoimmune
diseases. Macromolecular inhibitors of TNFα have proven therapeutically useful in some
preliminary studies. We have developed small molecule TNFα antagonist based on the
crystal structure of TNF receptor complex. The TNFα inhibitor is specific and mediates
biological function similar to the inhibitory soluble TNF receptor. This review focuses
on development of small molecule anti-TNFα mimetics by us and current status of other
agents.
INTRODUCTION Several biochemical and biological properties of TNFα
have been elucidated since the mid-1980s when TNFα was
cloned, sequenced and purified [4-7]. The major source of
TNFα is the activated monocytes/macrophages. TNFα is
synthesized as a 26 kDa soluble TNFα molecule and
observed as homotrimer under physiological conditions [8].
The main function of immune system is to eradicate
foreign organisms such as viruses or bacteria. Defense against
foreign organisms is mediated by innate immunity and by
specific (or adaptive) immunity. The effector phases of both
innate and specific immunity are large part mediated by
protein hormones called cytokines. In innate immunity, the
effector cytokines are mostly produced by mononuclear
phagocytes and are therefore called monokines. Phagocytes
accumulate at the site of infection and secrete monokines that
include interleukin-1 (IL-1), interleukin-6 (IL-6), interleukin-
8 (IL-8), interleukin-12 (IL-12) and tumor necrosis factor- α
(TNF α ). Most of these molecules are pleiotropic (i.e. affect
different biological functions) and have effects on
immunological processes such as inflammation and cellular
responses such as apoptosis.
Most of the cellular actions of TNFα have been
attributed to the activities of two distinct receptor molecules
TNF receptor I (TNFRI, p55) and TNFRII (p75) [9] which
are expressed ubiquitously. TNFRI-knock out mice are
resistant to endotoxin shock but succumb to infection,
indicating that TNFRI plays an important role in defense
against microorganisms [10]. The extracellular portions of
both TNF receptors can also be shed from the cell surface
through proteolytic cleavage and exist in soluble form and
moreover soluble receptors retain the ability to bind TNFα
and thus may act as physiological modulators of TNF
activity in vivo [11,12]. In the last century, Coley [1] observed beneficial
inflammatory effects in the terminally ill cancer patients.
Much later, in 1985, Old identified a protein in the serum of
endotoxin-treated rabbits that was responsible for the
hemorrhagic necrosis of tumors [2]. It was named tumor
necrosis factor (TNF) for its ability to trigger necrosis and
involution of transplantable tumors, and later named TNFα
after the discovery of lymphotoxin or TNFβ. TNFα was
highly toxic to both humans and animals [3]. In unrelated
experiments, cachectin isolated from waste body fluids of
animal and human with chronic disease were found to be
identical to the necrosis factor. Finally the study of
lipopolysaccharide (LPS) induced biological functions led to
the conclusion that TNFα is mediator of the shock,
disseminated coagulation, metabolic acidosis and end-organ
damage brought about by LPS.
Binding of TNFα to its membrane-bound receptors
induces diverse effects in different organs and tissues.
Recently several TNF receptor-associated proteins have been
cloned. The cytoplasmic domains of TNF receptors do not
have any intrinsic enzymatic activity, and hence they signal
by inducing aggregation of intracellular adaptor molecules.
The cytoplasmic domains of TNFRI bear a motif termed as
‘death domain’ (DD). The DD is a protein-protein
interaction motif that allows two proteins with DD to bind
to each other. Binding of TNF to TNFRI induces
recruitment of the DD-containing protein TRADD to the DD
of TNFRI [13]. Overexpression of TRADD also induces
TNF-regulated responses apoptosis and activation of the
transcription factors NF-κB and Jun kinase [14]. The group
of TNF receptor-associated factors (TRAF) also interact with
members of the TNFR family [15]. Most of TRAF proteins
interact with receptor molecules either directly, or indirectly
through binding to other TRAF, or through binding to
TRADD. TNFRII contains cytoplasmic TRAF binding
motifs and is able to bind directly to TRAF proteins.
Because TRAF2 can bind to TRADD, which in turn can
*Address correspondence to this author at the Department of Pathology
and Laboratory Medicine, 252 John Morgan Building, University of
Pennsylvania School of Medicine, 36th and Hamilton Walk, Philadelphia,
PA 19104, U.S.A; Tel: 215-573-9256; Fax: 215-898-2401; E-mail:
murali@xray.med.upenn.edu
** Current Address: 2-58 Hiromachi, 1-Chome, Shinagawa-Ku, Tokyo
140, Japan.
1389-5575/00 $20.00+.00 © 2001 Bentham Science Publishers, Ltd.