Retinal development in Drosophila: specifying the first neuron.

In vertebrates, a proneural basic helix-loop-helix transcription factor (Ath5, Atonal homolog 5) plays a crucial role in the specification of the first retinal neuron: the retinal ganglion cell (RGC). Math5 homozygous null mutant mice lack RGCs and have no optic nerve. Furthermore, the expression of the Ath5 protein is regulated to give a non-random dispersed pattern of RGCs. In Drosophila, retinal histogenesis is precisely coordinated and is associated with a progressive wave called the morphogenetic furrow. In the furrow, single precisely spaced cells are specified to become the first retinal neural cell type: the R8 photoreceptor cell. This Drosophila founder cell specification is coincident with and dependant upon the expression of the fly Ath5 ortholog: Atonal. Indeed, in both taxa, the process of founder cell specification may be viewed as the regulation of Atonal expression. It is now clear that, in flies, this regulation depends on the action of inductive and inhibitory signals. This review concentrates on the signaling mechanisms that produce this precise pattern of founder cells.