Patterns of importin-α expression during Drosophila spermatogenesis

Importin-α proteins do not only mediate the nuclear import of karyophilic proteins but also regulate spindle assembly during mitosis and the assembly of ring canals during Drosophila oogenesis. Three importin-α genes are present in the genome of Drosophila. To gain further insights into their function we analysed their expression during spermatogenesis by using antibodies raised against each of the three Importin-α proteins identified in Drosophila, namely, Imp-α1, -α2, and -α3. We found that each Imp-α is expressed during a specific and limited period of spermatogenesis. Strong expression of Imp-α2 takes place in spermatogonial cells, persists in spermatocytes, and lasts up to the completion of meiosis. In growing spermatocytes, the intracellular localisation of Imp-α2 appears to be dependent upon the rate of cell growth. In pupal testes Imp-α2 is essentially present in the spermatocyte nucleus but is localised in the cytoplasm of spermatocytes from adult testes. Both Imp-α1 and -α3 expression initiates at the beginning of meiosis and ends during spermatid differentiation. Imp-α1 expression extends up to the onset of the elongation phase, whereas that of Imp-α3 persists up to the completion of nuclear condensation when the spermatids become individualised. During meiosis Imp-α1 and -α3 are dispersed in the karyoplasm where they are partially associated with the nuclear spindle, albeit not with the asters. At telophase they aggregate around the chromatin. During sperm head differentiation, both Imp-α1 and -α3 are nuclear. These data indicate that each Imp-α protein carries during Drosophila spermatogenesis distinct, albeit overlapping, functions that may involve nuclear import of proteins, microtubule organisation, and other yet unknown processes.