Two new roles for the Drosophila AP patterning system in early morphogenesis.

Although gastrulation is regarded as the stage during Drosophila development when the AP patterning system first influences morphological processes, transcription is regulated in complex patterns already at cycle 10. How soon this transcriptional complexity produces spatial differences in morphology has been unclear. We report on two new processes that establish visible morphological inhomogeneities before the onset of gastrulation. The first of these is the regulation of syncytial nuclear densities in the anterior end of the egg and represents the first zygotically driven AP asymmetry in the embryo. The second process is the generation of a fine-scale pattern in the actin/myosin array during cellularization. We find three domains of different yolk stalk diameters as well as depths of cellularization along the AP axis. These domains are established under the control of the AP patterning system and require bicoid activity. The anterior-most domain is a region of large yolk stalk diameters and corresponds to the region of decreased nuclear densities observed during syncytial stages. The middle domain shows smaller yolk stalk diameters and more rapid cellularization. Its establishment requires wild-type paired activity and thus indirectly requires bicoid. It occurs in a region of the embryo that ultimately gives rise to the cephalic furrow and may account for the effect of paired on that structure during gastrulation. Our results therefore suggest a link between cytoskeletal organization during cellularization and subsequent morphogenetic processes of gastrulation.