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1
*
Department of Molecular Medicine, Osaka University Medical School, and
Osaka University, Suita City, Osaka, Japan
STAT-induced STAT inhibitor-1 (SSI-1), also referred to as
suppressor of cytokine signaling-1 and JAK-binding protein, is a member
of a new family, the members of which are negative regulators of
cytokine signals. SSI-1 is induced by various cytokines; however, the
transcriptional mechanism of the SSI-1 gene is not fully understood.
Here, we showed that transcription of the mouse SSI-1 gene was
initiated from six adjoining sites accompanying three GC boxes and a
single GC box-like element near them, but not from the TATA box or an
initiator sequence. We also showed that IFN-
induced SSI-1 mRNA more
strongly than IL-6 in NIH-3T3 fibroblasts and that this IFN- effect
was mediated by Stat1. To determine the signal pathway downstream of
Stat1, transcriptional activities of several mutant promoters were
examined. The region mediating stimulatory effect of IFN- to the
gene transcription was localized to the -88/-60 region containing
three tandem GAAA units, named variant IFN--responsive element
(VIRE), while four IFN- activation site (GAS)-like elements located
far upstream were not related to the IFN- response. Gel-shift assays
revealed that IFN- induced IFN regulatory factor-1 (IRF-1) binding
to VIRE, but not that of IRF-2 or three components of ISGF3.
Furthermore, forced expression of IRF-1 mimicked and that of IRF-2
inhibited the stimulatory effect of IFN- on SSI-1 gene
transcription. Finally, mouse embryonal fibroblasts lacking IRF-1
showed impaired SSI-1 mRNA induction by IFN-. These results
demonstrated that IRF-1, which is induced by activation of Stat1,
mediated transcriptional activation of the SSI-1 gene by IFN- via
VIRE.
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