We report the identification of a Drosophila Pax gene, eye gone (eyg), which is required for eye development. Loss-offunction eyg mutations cause reduction or absence of the eye. Similar to the Pax6 eyeless (ey) gene, ectopic expression of eyg induces extra eye formation, but at sites different from those induced by ey. Several lines of evidence suggest that eyg and ey act cooperatively: (1) eyg expression is not regulated by ey, nor does it regulate ey expression, (2) eyginduced ectopic morphogenetic furrow formation does not require ey, nor does ey-induced ectopic eye production require eyg, (3) eyg and ey can partially substitute for the function of the other, and (4) coexpression of eyg and ey has a synergistic enhancement of ectopic eye formation. Our results also show that eyg has two major functions: to promote cell proliferation in the eye disc and to promote eye development through suppression of wg transcription.

What are the developmental mechanisms required for conversion of an undifferentiated, two-dimensional field of cells into a patterned, tubular organ? In this report, we describe the contribution of the Drosophila Pax gene eye gone to the development of the embryonic salivary glands and ducts. eye gone expression in salivary tissues is controlled by several known regulators of salivary fate. After the initial establishment of the salivary primordium by Sex combs reduced, fork head excludes eye gone expression from the pregland cells so that its salivary expression is restricted to the posterior preduct cells. trachealess, in contrast, activates eye gone expression in the posterior preduct cells. We have previously described the process by which fork head and the EGF receptor pathway define the border between the gland and duct primordia. Here we show that eye gone is required for the subdivision of the duct primordium itself into the posterior individual duct and the anterior common duct domains. In the absence of eye gone, individual ducts as well as the precursor of the adult salivary glands, the imaginal ring, are absent. We took advantage of this ductless phenotype to show that Drosophila larvae do not have an obligate requirement for salivary glands and ducts. In addition to its role in the salivary duct, eye gone is required in the embryo for the development of the eye-antennal imaginal disc and the chemosensory antennal organ.

Organogenesis in Drosophila embryos begins at 4–5 hours of development as the expression of organ-specific genes is initiated. The salivary primordium, which occupies the ventral epidermis of parasegment 2, is among the earliest to be defined. It is soon divided into two distinct regions: the more dorsal pregland cells and the more ventral preduct cells. We show that it is the opposing activities of the Drosophila EGF receptor (DER) signaling pathway and the Fork head transcription factor that distinguish these cell types and set up the boundary between them. DER signaling acts ventrally to block fork head expression in the preduct cells, thereby restricting gland identity to the more dorsal cells. Fork head in turn blocks expression of duct-specific genes in the pregland cells, thereby restricting duct identity to the more ventral cells. A third regulatory activity, the Trachealess transcription factor, is also required to establish the identity of the preduct cells, but we show that it acts independently or downstream from the DER:fork head confrontation. In trachealess mutants, subdivision of the salivary primordium occurs normally and the dorsal cells form glands, but the ventral cells are undetermined. We present a model proposing that trachealess is the crucial duct-specific gene that Fork head represses to distinguish pregland from preduct cells.