Marbles mutants: uncoupling cell determination and nuclear migration in the developing Drosophila eye.

Morphogenesis of a multicellular structure requires not only that cells are specified to express particular gene products, but also that cells move to adopt characteristic shapes and positions. Little is known about how these two aspects of morphogenesis are coordinated. The developing Drosophila compound eye is a monolayer, in which cells are suspended between apical and basal membranes and assemble sequentially into hundreds of unit eyes, or facets, guided by a series of cell interactions. As cells are determined to join the facet, their nuclei and cell bodies rise apically and then settle into position in the cell group. The final nuclear positions determine the shape of the individual cells. We have identified a Drosophila gene called marbles which is required for the apical nuclear migrations that accompany cell determination during eye development. In marbles mutant eyes, the sequence of cell specification that leads to the formation of facets occurs almost normally despite the failure of nuclear migration in many cells. The marbles mutant phenotype reveals that during Drosophila eye development cell determination does not require nuclear migration.