Current Enzyme Inhibition, 2009, 5, 209-222 209
1573-4080/09 $55.00+.00 © 2009 Bentham Science Publishers Ltd.
Inhibition of Flaviviridae Replication Complex: Assays to Investigate the
Modulating Effect of Potential Compounds
Andrea Baier*
Department of Molecular Biology, John Paul II Catholic University of Lublin, Lublin, Poland
Abstract: Approximately up to 200 million new cases of infections caused by viruses of the Flaviviridae family, like
hepatitis C virus (HCV), West-Nile virus (WNV), dengue virus (DENV) and Japanese encephalitis virus (JEV) are regis-
tered annually.
Up to date, there is no effective antiviral therapy directed against Flaviviridae viruses. For the replication of all viruses the
intact function of nonstructural proteins is necessary. The blockade of their activities leads to inhibition of the virus
propagation. Thus, the nucleotide triphosphatase (NTPase) as well as helicase activities of NS3 and the RNA-dependent
RNA polymerase (RdRp) activity of NS5 (or NS5B) appear to be exceptionally attractive targets for antiviral compounds.
In this context, a systematic screening of chemical libraries and computational analysis are necessary which will lead to
the development of effective inhibitors of the Flaviviridae replication complex.
In this review the most promissing methods based on different techniques to quantify the enzymatic activities, like radio-
activity, fluorescence, colorimetry as well as a coupled second enzymatic reaction are summarized and discussed. These
methods serve as a starting point to establish a high throughput system (HTS) that permits more efficient screenings.
Keywords: Flaviviridae, in vitro assays, NTPase/helicase, RNA-dependent RNA polymerase.
The family Flaviviridae includes human and animal
pathogenic viruses of global importance e.g. the human
flaviviruses West-Nile virus (WNV), dengue virus (DENV),
Japanese encephalitis virus (JEV), tick-borne encephalitis
virus (TBEV) and yellow fever virus (YFV). This genus
consists of more than 70 species that are, on the basis of
phylogenetic analyses, divided into 14 classes which in turn
are grouped into three clusters: the mosquito-borne cluster,
the tick-borne cluster and the non-vector cluster [1]. All
flaviviruses of human importance are mosquito- or tick-
borne viruses which may cause fulminant, hemorrhagic dis-
eases and viral encephalitis. The second human pathogenic
genus containing hepatitis C virus (HCV) and GBV-B is
called hepacivirus and causes chronic or acute hepatits which
may lead to liver cirrhosis and hepatocellular carcinoma [2-
4]. Bovine viral diarrhea virus (BVDV), classical swine fe-
ver virus (CSFV) and border disease virus (BDV) [5] are
animal pathogenic viruses belonging to the genus pestivirus
and lead to severe diseases of the respective host usually
followed by death. Further viruses which are not classified in
one of the mentioned genera are GBV-A and GBV-C.
Possible routes of transmission differ between the genera.
HCV carriers are often intravenous-drug users or blood
product recipients. The latter could be reduced by blood
screening for HCV. Contrary to blood and sexual transmitted
HCV, flaviviruses are transmitted by vectors. Only rats and
mice are infected with no-vector viruses. In case of pestivi-
ruses transmission is directly from one animal to another by
the oro-nasal and transplacenteral route.
Flaviviridae viruses are small (40-50 nm) enveloped par-
ticles [2, 3, 6] with positive-sense single-stranded RNA
*Address correspondence to this author at the Department of Molecular
Biology, John Paul II Catholic University of Lublin, Al. Krasnicka 102, 20-
718 Lublin, Poland; Tel: +48 81 445 4603; Fax: +48 81 445 4611;
E-mail: baier@kul.lublin.pl
genome of the length of 9100 to 11000 bases which possess
a single open reading frame (ORF) flanked by 5‘- and 3‘-
terminal located untranslated regions (5’UTR and 3’UTR).
In the case of flaviviruses the 5‘UTR is relatively short (ap-
proximately 120 bases) and contains a cap structure type I
(m7GppN1mpN2). In hepaci- and pestiviruses the 5‘UTRs
consist of approximately 340 bases and reveal features of an
internal ribosomal entry site (IRES) mediating the binding to
the ribosome [7, 8]. There are some indications that the
adenosine- and/or uridine-rich stretches of the 3‘UTR are
involved in the regulation of the nucleoside triphosphatase
(NTPase) activity of the nonstructural protein 3 (NS3 pro-
tein) associated NTPase/helicase [9, 10]. Furthermore, it may
be important for the initiation of minus strand synthesis [11].
VIRAL REPLICATION
Genomic replication proceeds very similarly within in the
Flaviviridae family. After binding to the cell by specific re-
ceptors and entry of the virus via receptor-mediated endocy-
tosis the viral RNA is uncoated by acid-catalyzed membrane
fusion and translation is initiated by IRES- or cap-mediated
processes. The viral RNA is translated into one viral poly-
protein which is co- and posttranslationally cleaved by viral
and host proteases [12-14]. RNA synthesis occurs in cyto-
plasmic replication complexes localized at the perinuclear
membranes [15, 16]. All NS proteins appear to be involved
in RNA replication. The genomic RNA is encapsulated into
the core shell consisting of capsid proteins, enveloped by
viral surface protein embedded cellular derived lipid mem-
branes (Assembly).
The virus particle is budded through intracellular mem-
branes into cytoplasmic vesicles [17, 18]. By the cellular
secretory pathway the virus particles are transported to the
plasma membrane and undergo a maturation procedure. Vi-
rus release into the extracellular compartment is connected