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J Biol Chem, Vol. 273, Issue 27, 17206-17215, July 3, 1998

Regulation of Expression within a Gene Family
THE CASE OF THE RAT B- AND D-CRYSTALLIN PROMOTERS

From the Department of Molecular Biology, University of Nijmegen, Toernooiveld 1, 6525 ED Nijmegen, The Netherlands

The six closely related and clustered rat -crystallin genes, the A- to F-crystallin genes, are simultaneously activated in the embryonic lens but differentially shut down during postnatal development with the B-crystallin gene, the last one to be active. We show here that developmental silencing of the D-crystallin promoter correlates with delayed demethylation during lens fiber cell differentiation. Methylation silencing of the D-crystallin promoter is a general effect and does not require the methylation of a specific CpG, nor does methylation interfere with factor binding to the proximal activator. In later development, the D-crystallin promoter is also shut down earlier by a repressor that footprints to the 91/78 region. A factor with identical properties is present in brain. Hence, a ubiquitous factor has been recruited as a developmental regulator by the lens. All -crystallin promoters tested contain upstream silencers, but at least the B-crystallin silencer is distinct from the D-crystallin silencer. The -crystallin promoters were found to share a proximal activator (the -box; around 50), which behaves as a MARE. The B-box is recognized with much lower avidity than the D-box. By swapping elements between the B- and the D-crystallin promoter, we show that activation by the B-box requires a directly adjacent 46/38 AP-1 consensus site. These experiments also uncovered another positive element in the D-crystallin promoter, around 10. In the context of the D-crystallin promoter, this element is redundant; in the context of the B-crystallin promoter, it can replace the 46/38 element.

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