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J Biol Chem, Vol. 275, Issue 15, 10761-10766, April 14, 2000
From the Departments of In our previous study we showed that
insulin-like growth factor-I induces a cAMP-response element (CRE)
site-containing Bcl-2 promoter through a novel signaling pathway
involving mitogen-activated protein kinase kinase 6/p38
Akt/Protein Kinase B Up-regulates Bcl-2 Expression through
cAMP-response Element-binding Protein*
§,§,
,§
Endocrinology,
¶ Pharmacology, and ** Medicine, University of Colorado Health
Sciences Center, Denver, Colorado 80262, the § Section of
Endocrinology, Veterans Affairs Medical Center, Denver, Colorado 80220, and Department of Medicine, Stanford University School of
Medicine, Stanford, California 94305
mitogen-activated protein kinase/MAP kinase-activated protein
kinase-3/cAMP-response element-binding protein (CREB)
(Pugazhenthi, S., Miller, E., Sable, C., Young, P., Heidenreich,
K. A., Boxer, L. M., and Reusch, J. E.-B. (1999) J. Biol. Chem. 274, 27529-27535). In the present
investigation, we define a second pathway contributing to
CREB-dependent up-regulation of Bcl-2 expression as a novel
anti-apoptotic function of Akt signaling. To examine the role of Akt on
Bcl-2 expression, a series of transient transfections using a
luciferase reporter gene driven by the promoter region of Bcl-2
containing a CRE were carried out. Pharmacological inhibition of
phosphatidylinositol (PI) 3-kinase, the upstream kinase of Akt, with
LY294002 led to a 45% decrease in Bcl-2 promoter activity. The
reporter activity was enhanced 2.3-fold by overexpression of active
p110 subunit of PI 3-kinase and inhibited 44% by the dominant negative
p85 subunit of PI 3-kinase. Cotransfection with
3-phosphoinositide-dependent kinase (PDK1), which is required
for the full activation of Akt, resulted in enhanced luciferase
activity. Insulin-like growth factor-I-mediated induction of Bcl-2
promoter activity was decreased significantly (p < 0.01) by the dominant negative forms of p85 subunit of PI 3-kinase,
PDK1, and Akt. These data indicate that regulation of Bcl-2 expression
by IGF-I involves a signaling cascade mediated by PI
3-kinase/PDK1/Akt/CREB. Furthermore, we measured the Bcl-2 mRNA in
PC12 cells overexpressing Akt by real-time quantitative reverse
transcription-polymerase chain reaction using the TaqManTM
fluorogenic probe system. We observed a 2.1-fold increase in Bcl-2
mRNA levels in the Akt cell line compared with control PC12 cells,
supporting the observation that enhanced CREB activity by Akt signaling
leads to increased Bcl-2 promoter activity and cell survival.
*
The work was supported by a Veterans Affairs Merit review
grant and an American Diabetes Association research award and National Institute of Health Grant KO8 DK02351 (NIDDK) (to J. E.-B. R.) and a
Veterans Affairs Research Enhancement Award Program grant (to
J. E.-B. R. and K. A. H.).The costs of publication of this article were defrayed in part by the
payment of page charges. The article
must therefore be hereby marked
"advertisement" in accordance with 18 U.S.C. Section
1734 solely to indicate this fact.
To whom correspondence should be addressed: Section of
Endocrinology (111H), Veterans Affairs Medical Center, 1055 Clermont St., Denver, CO 80220. Tel.: 303-399-8020 (ext. 2775); Fax:
303-393-5271; E-mail: ReuschJ@den-res.org.
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