The imaginal discs of Drosophila are a useful experimental system in which we can study the origin and genetic determination of spatial patterns in development. This involves the separation of the disc-cell population into distinct lineage compartments, based on clonally transmitted expression states of a number of known selector genes. However, these commitments can be abrogated and the compartment boundaries redeployed, when repatterning occurs in cultured disc fragments. This has so far only been explained using the idea of positional information. The genetic basis of this property of the imaginal disc system and its relationship to compartments have not been identified. Here we have screened over 470 recessive lethal P-lacZ enhancer-trap insertions from the Berkeley Drosophila Genome Project for expression after cell death, which initiates pattern respecification in the imaginal discs. The positive lines obtained identify essential genes that may be important for pattern formation. Most show patterned imaginal disc expression, and many have maternal or zygotic effects on embryonic development. One is an allele of schnurri, a gene that encodes a component of the decapentaplegic (dpp) signal transduction pathway used for positional signalling in the embryo and in imaginal discs.
When imaginal disc fragments from Drosophila are cultured in adult female hosts, they either duplicate the part of the pattern specified by the fate map, or regenerate to replace the missing part. The new tissue is added by proliferation of a small number of cells from the cut edge, brought together when the wound heals to form a regeneration blastema. Specification of the new pattern has been explained by assuming interactions among cells of different positional value in the regeneration blastema. In order to identify genes which might mediate these events, we screened over eight hundred independently isolated autosomal insertions of an enhancer-sensitive P-element, for altered lac-z expression in regenerating discs following cell death induced by a temperature-sensitive cell-lethal mutation. Two further screens divided the positive lines into four groups based on appropriate timing of the lac-z response in the cell-lethal mutant background and the expected response to an alternate source of cell death. Expression in wing disc fragments cultured in vivo was most frequent in the target class defined by the screens. In this direct test, lac-z expression was found in 23 lines and in most cases was spatially and temporally correlated with the formation of the regeneration blastema. Our results suggest a very substantial transcriptional response during the early stages of imaginal disc regeneration. lac-z expression in control imaginal discs, embryos and adult ovaries of the positive lines was also assayed. The selected insertions included: a small class expressed only in discs undergoing regeneration and apparently not at any other stage, possibly representing genes active exclusively in regeneration; a larger class expressed in the embryo or during oogenesis, but not normally in imaginal discs, as expected for functions recruited from earlier stages of the developmental program; and finally a class with spatially patterned expression in normal discs. This class included several insertions with expression associated with compartment boundaries, including one at the decapentaplegic (dpp), and one at the crumbs (crb) locus, a growth factor homologue, and an EGF-repeat gene respectively. Some of the expression patterns observed in cultured disc fragments provide evidence for cell communication in the regeneration blastema.
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