Dorsoventral pattern of Drosophila embryo is specified by the nuclear localization gradient of the transcription factor Dorsal. Genetic and genomic studies of this morphogen gradient provided important insights into spatial control of gene expression in development. Recent live imaging experiments revealed hitherto unappreciated dynamics of the Dorsal gradient and posed new questions about the mechanisms of its transcriptional interpretation. Some of these questions can be answered by models in which the morphogenetic capacity of the Dorsal gradient is potentiated by spatially uniform factors, such as Zelda, a transcription factor that plays a key role in the activation of zygotic transcription. Combinatorial effects of uniform and graded factors play an important role in the transcriptional and signaling cascades initiated by Dorsal and may explain differential positioning of gene expression borders by other morphogen gradients.
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