Reshaping the Syncytial Drosophila Embryo with Cortical Actin Networks: Four Main Steps of Early Development.

Drosophila development begins as a syncytium. The large size of the one-cell embryo makes it ideal for studying the structure, regulation, and effects of the cortical actin cytoskeleton. We review four main steps of early development that depend on the actin cortex. At each step, dynamic remodelling of the cortex has specific effects on nuclei within the syncytium. During axial expansion, a cortical actomyosin network assembles and disassembles with the cell cycle, generating cytoplasmic flows that evenly distribute nuclei along the ovoid cell. When nuclei move to the cell periphery, they seed Arp2/3-based actin caps which grow into an array of dome-like compartments that house the nuclei as they divide at the cell cortex. To separate germline nuclei from the soma, posterior germ plasm induces full cleavage of mono-nucleated primordial germ cells from the syncytium. Finally, zygotic gene expression triggers formation of the blastoderm epithelium via cellularization and simultaneous division of ~6000 mono-nucleated cells from a single internal yolk cell. During these steps, the cortex is regulated in space and time, gains domain and sub-domain structure, and undergoes mesoscale interactions that lay a structural foundation of animal development.