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J Biol Chem, Vol. 274, Issue 49, 35037-35045, December 3, 1999
From the Centre de Recherche en Reproduction Animale, Faculté
de Médecine Vétérinaire, Université de
Montréal, C.P. 5000, Saint-Hyacinthe,
Québec J2S 7C6, Canada
To elucidate the molecular
mechanisms involved in the delayed induction of PGHS-2 in species with
a long ovulatory process, a 1.6-kilobase fragment of the bovine PGHS-2
promoter was isolated, and its activity was characterized in primary
cultures of bovine granulosa cells. Promoter activity assays performed
with a series of deletion mutants revealed that the promoter region
from 
149 to 2 (+1 = transcription start site) confers
full-length promoter activity in response to forskolin (10 µM). Four consensus cis-elements were
identified within this region, including an E-box, ATF/CRE, C/EBP, and
AP2 site. Site-directed mutagenesis showed that the E-box was required
for PGHS-2 promoter activity, that disruption of the C/EBP element
decreased forskolin inducible activity by 29%, whereas point mutation
within the ATF/CRE and AP2 element had no inhibitory effect.
Electrophoretic mobility shift assays (EMSAs) performed with the
149/2 fragment and granulosa cell nuclear extracts obtained before
(0 h) and after (18 and 20 h) human chorionic gonadotropin (hCG)
revealed the regulation of multiple DNA-protein complexes. The 0-h
extract generated four complexes at the E-box, whereas only one complex
was produced at this site with the 18-h extract. Supershift EMSAs
identified that upstream stimulatory factor-1 and -2 (USF-1 and -2)
were part of these complexes. Interestingly, the presence of the
amino-terminal truncated USF-2, which lacks the transcription
activation domain, was detected in the 0-h extract, but not in extracts
prepared post-hCG. Supershift EMSAs also indicated high levels of
C/EBP
binding to its cis-element in the 0-h extract,
which contrasts with results previously reported in rats. Thus, high
levels of amino-terminal truncated USF-2 and C/EBP in bovine
granulosa cells prior to hCG treatment could repress gene expression,
and be involved in the delayed induction of PGHS-2 in species with a
long ovulatory process.
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