15 E.J. Schaefer (ed.), High Density Lipoproteins, Dyslipidemia, and Coronary Heart Disease,
DOI 10.1007/978-1-4419-1059-2_2, © Springer Science+Business Media, LLC 2010
Abbreviations
C/EBP CAAT/enhancer binding protein
CAT Chloramphenicol acetyl transferase
EGR-1 Early growth response factor-1
FXR Farnesoid X receptor
HNF-4 Hepatocyte nuclear factor-4
HDL High density lipoprotein
HRE Hormone response element
LRH-1 Liver receptor homolog-1
LXRs Liver X receptors
PPARa Peroxisome proliferator-activated receptor a
PLTP Phospholipid transfer protein
RORa Retinoic acid receptor-related orphan receptor a
RXRa Retinoid X receptor a
SHP Small heterodimer partner
SP1 Specificity protein 1
SR-BI Scavenger receptor class B type I
SREBP Sterol regulatory element binding protein
WT Wild type
Transcriptional Regulation
of the Human ApoA-I Gene in Cell Culture
and in ApoA-I Transgenic Mice
Role of hormone nuclear receptors, SP1 and of a common
enhancer on the transcriptional regulation of the human
apoA-I gene in cell cultures. Earlier studies established that
there is a linkage and common regulatory mechanism of the
apoA-I/apoCIII/apoA-IV gene cluster (Fig. 1a). In this cluster,
the distal regulatory region of the apoCIII promoter is an
enhancer that increases the strength of the neighboring
promoters in vitro [1–4]. When the apoCIII enhancer is
joined with the proximal apoA-I promoter, then the activity
of the promoter increases over tenfold [1–4]. The proximal
apoA-I promoter contains two hormone response elements
(HREs) that bind orphan and ligand-dependent nuclear
receptors [5, 6]. The apoCIII enhancer also has two HREs
which differ in their receptor specificity, and also has three
SP1 binding sites [1, 7] (Fig. 1a).
Systematic in vitro mutagenesis of different sites of the
promoter/enhancer cluster and determination of the promoter
activity by CAT assays showed that mutations in the HRE,
and the SP1 binding sites affect the activity of the apoA-I
promoter/enhancer cluster in HepG2 cells [1–7]. Mutation in
the proximal HREs reduced the strength of the promoter/
enhancer to 6% of the WT control, whereas mutations in the
HRE (I
4
) of the enhancer reduced the activity of the promoter
enhancer cluster to 18% of the WT control (Fig. 1b). Finally,
the individual mutations in the SP1 binding site reduced the
activity of the promoter enhancer cluster to 40–60% of the
WT control (Fig. 1b).
The receptor specificity of the HREs of the apoA-I promoter
and the apoCIII enhancer were determined by DNA binding
gel electrophoresis assays. These analyses established that
both HREs present in the proximal apoA-I promoter bind
HNF-4, other orphan receptors, and a variety of ligand-
dependent nuclear receptors with different affinities [6]
(Fig. 1c). One of the HREs of the enhancer binds also HNF-4,
other orphan nuclear receptors, and ligand-dependent nuclear
receptors with different affinities, whereas the other HRE
does not bind HNF-4, binds other orphan receptors, and
different combinations of ligand-dependent nuclear receptors
with different affinities [7] (Fig. 1c).
Transcriptional regulation of the human apoA-I gene in
transgenic mice. To validate the conclusions drawn by the
in vitro experiments, we generated a variety of transgenic
mouse lines which express the WT A-I/CIII cluster or the
same cluster with the mutations in the HREs and the binding
sites of SP1 and other transcription factors. In these constructs,
the apoCIII gene was replaced by the CAT gene (Fig. 2).
V.I. Zannis (*)
Whitaker Cardiovascular Institute,
Boston University School of Medicine,
Boston, MA, USA
e-mail: vzannis@bu.edu
Regulation of ApoA-I Gene Expression and Prospects
to Increase Plasma ApoA-I and HDL Levels
Vassilis I. Zannis, Adelina Duka, Konstantinos Drosatos, Despina Sanoudou,
Georgios Koukos, Eleni Zanni, and Dimitris Kardassis