Inhibition of Cyclic AMP Response Element-Binding
Protein/Cyclic AMP Response Element-Mediated Transcription
by the Immunosuppressive Drugs Cyclosporin A and FK506
Depends on the Promoter Context
GERO SIEMANN, ROLAND BLUME, DANIELA GRAPENTIN, ELKE OETJEN, MARKUS SCHWANINGER
1
, and
WILLHART KNEPEL
Department of Molecular Pharmacology, University of Go ¨ ttingen, Go ¨ ttingen, Germany
Received October 26, 1998; accepted February 22, 1999 This paper is available online at http://www.molpharm.org
ABSTRACT
The immunosuppressants cyclosporin A and FK506 (tacroli-
mus) can block the phosphatase calcineurin, thereby inhibiting
gene transcription directed by the cyclic AMP (cAMP)- and
calcium-responsive transcription factor, cAMP response ele-
ment (CRE)-binding protein, and its binding site, CRE, in vari-
ous cell lines. This action is a novel molecular mechanism of
cyclosporin A and FK506 action. Because inhibition of CREB/
CRE-directed transcription by cyclosporin A and FK506 has
previously been observed by using synthetic minienhancers,
reporter fusion genes were constructed to examine the effect of
cyclosporin A and FK506 on the transcriptional activity of CRE-
containing natural promoters. In transient transfection experi-
ments, cyclosporin A and FK506 inhibited the transcriptional
activation by cAMP and the membrane depolarization of three
CRE-containing promoters. However, cyclosporin A and FK506
failed to inhibit the activation by cAMP of another promoter, the
rat insulin I gene promoter. The lack of cyclosporin A/FK506
sensitivity is not intrinsic to the insulin CRE because cyclo-
sporin A and FK506 inhibited the activation by cAMP of the
insulin CRE when isolated and used as a synthetic minien-
hancer. Rather, cyclosporin A/FK506 resistance may be con-
ferred by specific promoter interactions because a mutational
analysis of the insulin promoter revealed that inside this pro-
moter, CRE activity depends on an adjacent control element.
These data show that cyclosporin A and FK506 can inhibit CRE
activity when the CRE resides in its natural promoter. However,
the cyclosporin A/FK506 sensitivity depends on the specific
promoter context. The results suggest that cyclosporin A and
FK506 may alter target tissue function through the regulation of
a subset of CRE-containing genes.
Cyclosporin A and FK506 (tacrolimus) are clinically impor-
tant immunosuppressive drugs that are widely used to pre-
vent graft rejection after organ transplantation. The intro-
duction of cyclosporin A in the field of organ transplantation
in the early 1980s resulted in extraordinary improvements of
graft survival, and cyclosporin A has become a first-choice
drug for patients with allograft organs. In addition, cyclo-
sporin A is used in the therapy of an increasing number of
autoimmune diseases. However, the therapeutic application
of cyclosporin A and FK506 is limited by untoward effects
that are shared by both drugs, including nephrotoxicity, hy-
pertension, neurotoxicity, and impaired glucose tolerance
(European FK506 Multicenter Liver Study Group, 1994; U.S.
Multicenter FK506 Liver Study Group, 1994). The structur-
ally unrelated drugs bind to their respective intracellular
receptors, the immunophilins. These drug/immunophilin
complexes directly target the calcium/calmodulin-dependent
phosphatase calcineurin, thereby blocking its activity (Ho et
al., 1996). To date, all of the therapeutic effects, as well as the
toxic effects, of these drugs have been shown to be due to
inhibition of calcineurin. Inhibition of calcineurin blocks the
translocation of the cytosolic component of the nuclear factor
of activated T cells (NFAT) into the nucleus, resulting in a
failure to activate the genes regulated by the NFAT tran-
scription factor, including those necessary for T cell prolifer-
ation, such as interleukin 2 (Ho et al., 1996; Rao et al., 1997;
Ru ¨ hlmann and Nordheim, 1997). Inhibition of NFAT-di-
rected transcription may be important for the repression of
early steps in T cell activation and, thus, immunosuppression
induced by cyclosporin A and FK506 (Ho et al., 1996; Rao et
al., 1997; Ru ¨ hlmann and Nordheim, 1997). However, other
targets of calcineurin are likely to play a role in this process.
This work was supported by Deutsche Forschungsgemeinschaft Grant
SFB402/A3.
1
Present address: Department of Neurology, University of Heidelberg, Hei-
delberg, Germany.
ABBREVIATIONS: CAT, chloramphenicol acetyltransferase; CRE, cAMP response element; CREB, CRE-binding protein; NFAT, nuclear factor of
activated T cells; bp, base pair.
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