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Subunit Gene Expression
Department of Biochemistry, Molecular Biology, and Cell Biology
(A.M., K.E.M.) Northwestern University Evanston, Illinois
60208
Department of Physiology and Biophysics (J.U.)
Chicago Medical School North Chicago, Illinois 60064
Institut de Genetique et de Biologie Moleculaire et
Cellulaire (P.S.-C.) Centre National de la Recherche
Scientifique-INSERM-ULP BP163, 67404 Ilkirch, Strasbourg,
France
Many hormones that stimulate intracellular
signaling pathways utilizing the second messenger cAMP affect gene
expression in target cells through the activation of cAMP-responsive
transcriptional regulatory proteins. Two of the best characterized of
these are the cAMP-response element (CRE)-binding protein (CREB) and
the CRE-modulatory protein (CREM). CREB and CREM are expressed as a
family of proteins that have diverse activities in either stimulating
or repressing gene transcription. In this study we examined the
expression and regulation of the CREM gene in the rat ovary and in
granulosa cells, to determine whether repressor isoforms of CREM might
have a role in the LH-mediated suppression of inhibin 
-subunit gene
expression that occurs just before ovulation. We found that the
predominant CREM mRNAs in the ovary correspond to previously described
internal transcripts of the CREM gene that encode the inducible cAMP
early repressor (ICER). ICER mRNAs are strongly induced in the ovary by
exogenous gonadotropins in immature rats and are transiently expressed
in the ovary immediately after the preovulatory LH surge in adult
cycling rats. Although ICER is expressed in multiple ovarian cell
types, expression in granulosa cells is observed only in response to LH
stimulation. ICER mRNAs are also induced by the activation of
cAMP-signaling pathways in cultured primary granulosa cells. To
determine whether ICER can act as a functional repressor to modulate
potential target genes such as the inhibin -subunit gene, an ICER
expression construct was transiently cotransfected into a granulosa
cell line along with an inhibin -subunit promoter-luciferase
reporter gene. Both basal and cAMP-induced expression of the inhibin
-subunit promoter were suppressed by ICER. These studies reveal that
CREM, a tissue-specific factor, is expressed and regulated by
gonadotropins in the ovary, that the predominant CREM transcripts
encode the repressor protein ICER, and that ICER is capable of
inhibiting cAMP-induced expression of the inhibin -subunit gene. Our
findings are consistent with a role for repressors such as ICER in
mediating the suppression of inhibin -subunit gene expression that
occurs in the ovary at the time of the preovulatory LH surge.
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