Recent Patents on Anti-Infective Drug Discovery, 2006, 1, 67-73 67
1574-891X/06 $100.00+.00 © 2006 Bentham Science Publishers Ltd.
An Insight on the Leading HIV Entry Inhibitors
Ana Salomé Veiga
1
, Nuno C. Santos
2
and Miguel A. R. B. Castanho*
,1
1
Centro de Química e Bioquímica, Faculdade de Ciências da Universidade de Lisboa, Campo Grande Ed. C8, 1749-016
Lisboa, Portugal;
2
Instituto de Biopatologia Química / Faculdade de Medicina de Lisboa and Unidade de Biopatologia
Vascular / Instituto de Medicina Molecular, Av. Prof. Egas Moniz, 1649-028 Lisboa, Portugal
Received: July 03, 2005; Accepted: August 17, 2005; Revised: September 14, 2005
Abstract: The main strategies nowadays to fight AIDS rely on chemical therapy to inhibit the reverse transcriptase or
protease of HIV. However, a synthetic 36 amino-acids peptide that blocks the entry of the virus in the target cells
(enfuvirtide) has recently reached approval for clinical application. This molecule may probably be just the leader of a
new generation of drugs that is about to emerge to interrupt the first step in the HIV life cycle, i.e. preventing the virus
from actually entering cells. This paper reviews the enfuvirtide path from clinical trials to the attempts to detail its
molecular-level mode of action. It is commonly accepted that this peptide would block the fusion between viral and cell
plasma membrane through binding to the N-terminal heptad repeat (NHR) region of the viral protein gp41. However,
there has been growing evidence that this model of action may be unrealistic, the action of enfuvirtide being more
complex and diverse than initially thought. Membrane-assisted local concentration increase and interference with
gp120/co-receptor docking may also contribute for the inhibitory action of the peptide. Selected HIV-entry inhibitors on
clinical trials are presented to characterize the future drugs in the market in this class.
Keywords: Entry, attachment, fusion, binding, entrance inhibitor, HIV, AIDS, virus, chemokine, peptide, receptor, co-receptor,
synergism, clinical trial, enfuvirtide.
1. INTRODUCTION
Combination therapy with reverse transcriptase and
protease inhibitors is the most common current treatment of
HIV-1 infection [1,2]. Despite the success of this therapy,
namely reducing morbidity and mortality of HIV-1 infected
patients [3-6], it has adverse effects and drug resistant HIV-1
strains emerged [7-11]. A new class of antiviral agents is in
development, the entry inhibitors. These molecules target the
first step in the HIV-1 replication cycle, the viral entry [12-
15]. Unlike reverse transcriptase and protease inhibitors,
which target post-entry steps, entry or fusion inhibitors act
extracellularly preventing viral entry into target cells. The
most advanced and already approved by the Food and Drug
Administration fusion inhibitor is enfuvirtide (T20; DP-178)
from Trimeris/Roche [16].
HIV-1 Entry
The HIV-1 envelope glycoproteins complex is expressed
on the surface of the viral membrane as an oligomeric
protein (trimer) and mediates the viral entry. The complex is
composed of two subunits noncovalently associated: gp120,
the surface glycoprotein, which interacts with cellular
receptors, and gp41, the transmembrane glycoprotein,
responsible for fusion of the viral and cellular membranes
[e.g. 12,17,18]. HIV-1 entry into target cells is a multi-step
process that is initiated by gp120 binding to the CD4
receptor, present in the target cells surface. This contact
*Address correspondence to this author at the Centro de Química e
Bioquímica, Faculdade de Ciências, Universidade de Lisboa, Campo
Grande C8, 1749-016 Lisboa, Portugal; Tel: + 351 21 7500931; Fax: +351
21 7500088; E-mail: castanho@fc.ul.pt
induces a conformational change in gp120 enabling it to bind
to a cellular co-receptor, usually CCR5 or CXCR4. gp120
binding to CD4 and co-receptor induces conformational
changes in gp41 exposing the N-terminal fusion peptide and
allowing its insertion into target cell membrane, thereby
connecting both membranes. Subsequent changes within the
gp41 ectodomain involve the interaction of two heptad
repeat sequences (HR1, next the fusion peptide and HR2,
preceding the transmembrane domain) and a six-helix bundle
structure (also called hairpin structure) is formed. The
hairpin formation brings the viral and cell membrane into
closed proximity, allowing fusion of the membranes and then
entry of the virus [12,13,17-19].
Each of the main steps in the HIV-1 entry process can be
a target for entry inhibitors. The ones currently under
development fall into three categories: gp120-CD4 binding
inhibitors, gp120-co-receptors binding inhibitors, and
membrane fusion inhibitors, which interfere with gp41
conformational changes [2,14,16,20]. Fusion inhibitors are
the leading compounds, one of them, enfuvirtide, being
already approved for clinical applications.
2. ENFUVIRTIDE CLINICAL TRIALS
The publication in 1998 of the results of the TRI-001
clinical trial [21] is considered as the proof-of-concept that
the entrance of the virus in the cell of the host can be blocked
in vivo by enfuvirtide. This study evaluated the use of
enfuvirtide subcutaneous infusion, as monotherapy, during
14 days. The TRI-001 allowed concluding that the
administration, during a reduced period, is safe and induces a
significant (1.14 log
10
) viral charge decrease in the group
that received a 90 mg deliverable dose of enfuvirtide (100
mg nominal dose) twice daily.