Organization of the cytoskeleton in early Drosophila embryos

The cytoskeleton of early, non-cellularized Drosophila embryos has been examined by indirect immunofluorescence techniques, using whole mounts to visualize the cortical cytoplasm and sections to visualize the interior. Before the completion of outward nuclear migration at nuclear cycle 10, both actin filaments and microtubules are concentrated in a uniform surface layer a few micrometers deep, while a network of microtubules surrounds each of the nuclei in the embryo interior. These two filament-rich regions in the early embryo correspond to special regions of cytoplasm that tend to exclude cytoplasmic particles in light micrographs of histological sections. After the nuclei in the interior migrate to the cell surface and form the syncytial blastoderm, each nucleus is seen to be surrounded by its own domain of filament-rich cytoplasm, into which the cytoskeletal proteins of the original surface layer have presumably been incorporated. At interphase, the microtubules seem to be organized from the centrosome directly above each nucleus, extending to a depth of at least 40 microns throughout the cortical region of cytoplasm (the periplasm). During this stage of the cell cycle, there is also an actin "cap" underlying the plasma membrane immediately above each nucleus. As each nucleus enters mitosis, the centrosome splits and the microtubules are rearranged to form a mitotic spindle. The actin underlying the plasma membrane spreads out, and closely spaced adjacent spindles become separated by transient membrane furrows that are associated with a continuous actin filament-rich layer. Thus, each nucleus in the syncytial blastoderm is surrounded by its own individualized region of the cytoplasm, despite the fact that it shares a single cytoplasmic compartment with thousands of other nuclei.