The engrailed gene product of Drosophila specifies the fate of a subset of cells in each segment. Our studies of engrailed regulation suggest that fate determination is an elaborate, multistep process. At the time in embryogenesis when the engrailed-dependent cell fate is probably determined, four modes of control act in an overlapping progression to govern engrailed expression. After activation by pairrule genes, both an extracellular signal, wingless, and autoregulation are required for engrailed expression. Autoregulation graduates to wingless independence, but is transient, and is superseded by an engrailed-independent mode of maintenance.Two levels of commitment have been defined for the choice of cell fate 1 . Early in development, fate can be instructed by environmental cues. This fate specification is flexible and allows fate to change in response to extracellular signals. But cells can undergo a transition from this type of specification to one which is determined and therefore irreversible. Because a determined cell maintains a particular fate independent of its environment, the transition to a determined state requires that development comes under cell-intrinsic, rather than cell-extrinsic control. The identification of selector genes in Drosophila provides an opportunity to study the basis of both types of cell-fate control. Selector genes encode regulators whose function defines the fate of a cell, and whose continued expression stably maintains cell fate 2,3 . Where the choice of cell fate is initially dependent on the environment, selector gene expression must be sensitive to extracellular signals. By contrast, once cell fate is determined, expression of these genes must be governed exclusively by cell-intrinsic factors.The engrailed (en) gene is a selector gene that distinguishes two populations of cells in each developing embryonic segment, the progenitors of the anterior and posterior compartments 3,4 . This distinction can be seen as a lineage restriction when individual embryonic cells are marked by mitotic recombination 5,6 . These give rise to clonal patches in the adult that never span the boundary between the anterior and posterior compartments, demonstrating that, from the time of marking, the progeny of a given cell only contribute to one compartment. This restriction requires expression of en by posterior cells 3,4,7 . But it seems that the onset of localized en expression in early embryos is not sufficient to irreversibly determine posterior compartment identity because en requires at least one cell-extrinsic input at this time.This input is provided by the wingless (wg) gene, expressed in segmentally repeated stripes of cells that lie next to en-expressing cells 8,9 . The wg protein, which is homologous to the product of the proto-oncogene Wnt-1 ( ref. 10 ), is secreted and appears to be taken up by en-expressing cells 9 . In wg mutant embryos, en expression is initiated in a normal striped pattern but soon stops, indicating that continued en expression in wild-type embryos r...