Asymmetric localization of Drosophila pair-rule transcripts from displaced nuclei: evidence for directional nuclear export.

Drosophila pair-rule transcripts accumulate exclusively apical of the layer of peripheral nuclei in syncytial blastoderm stage embryos. Here, we use aneuploid embryos to test zygotic gene requirements for pair-rule transcript localization. As apical localization is maintained in all genotypes tested, the required components must be maternally encoded. In aneuploid embryos with multiple layers or cortical nuclei, pair-rule transcripts lie apical of both superficial and internalized nuclei. In the latter case, the transcripts are 'pseudo-apical', i.e. apical of the nuclei from which they derive, but basal of superficial nuclei. We show that internalized nuclei maintain their apico-basal nuclear orientation, and that they lack the apical cytoskeletal assemblies which lie adjacent to superficial nuclei. These results support a mechanism of localizing pair-rule transcripts by directional (vectorial) nuclear export.