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Vol. 15, No. 3, pp. 340-351, February 1, 2001
Department of Biology, New York University, New York, New York
10003, USA
The establishment of expression domains of developmentally regulated
genes depends on cues provided by different concentrations of
transcriptional activators and repressors. Here we analyze the
regulation of the Drosophila gene zen, which is a
target of the Decapentaplegic (Dpp) signaling pathway during cellular
blastoderm formation. We show that low levels of the Dpp signal
transducer p-Mad (phosphorylated Mad), together with the recently
discovered negative regulator Brinker (Brk), define the spatial limits
of zen transcription in a broad dorsal-on/ventral-off domain.
The subsequent refinement of this pattern to the dorsal-most cells, however, correlates with high levels of p-Mad that accumulate in the
same region during late blastoderm. Examination of the zen
regulatory sequences revealed the presence of multiple Mad and Brk
binding sites, and our results indicate that a full occupancy of the
Mad sites due to high concentrations of nuclear Mad is the primary
mechanism for refinement of zen. Interestingly, several Mad and
Brk binding sites overlap, and we show that Mad and Brk cannot bind
simultaneously to such sites. We propose a model whereby competition
between Mad and Brk determines spatially restricted domains of
expression of Dpp target genes.
[Key Words: Dpp morphogen; target genes; Smad activation; Brk repression]
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