Pattern formation in a secondary field: a hierarchy of regulatory genes subdivides the developing Drosophila wing disc into discrete subregions

The legs and wings of insects and vertebrates develop from secondary embryonic fields that arise after the primary body axes have been established. In order to understand how the insect imaginal wing field is patterned, we have examined in detail the temporal and spatial expression patterns of, and epistatic relationships between, four key regulatory genes that are specifically required for wing formation in Drosophila. The wingless protein, in a role surprisingly distinct from its embryonic segment polarity function, appears to be the earliest-acting member of the hierarchy and crucial for distinguishing the notum/wing subfields, and for the compartmentalization of the dorsal and ventral wing surfaces. The wingless product is required to restrict the expression of the apterous gene to dorsal cells and to promote the expression of the vestigial and scalloped genes that demarcate the wing primordia and act in concert to promote morphogenesis.