Ultrastructure of capitate projections in the optic neuropil of Diptera

Photoreceptor axons in the first optic neuropil of the dipteran flies Musca domestica and Drosophila melanogaster was examined with electron microscopy. The objective was to determine ultrastructure, persistence and glial source of the capitate projections found within these neurons. Capitate projections are simple or compound processes of epithelial glial cells which profusely insert into form-fitting folds of axon terminals of the peripheral retinular cells (R1-6) in the synaptic plexus portion of the first optic neuropil. These neuro-glial junctions may be simple indentations, have a head with a single stalk, or possess a single, circular stalk from which 3 or 4 bulbous (glial) heads are elaborated. Using serial thick sections of Drosophila neuropil for HVEM we were able to observe that the stalks connecting nearly all capitate projections led directly to a glial cell. Thus no disembodied heads were found suspended in axoplasm. Capitate projections appeared to be persistent structures, present in young as well as senescent adults. No evolution of form was found; thus 3 distinct expressions of these glial processes (without transitional forms) are present. From freeze-fracture replicas and serial HVEM sections it was determined that there were approximately 3 capitate projections per micron 2 in Drosophila and Musca, respectively. About 800 capitate projections exist per Musca axon terminal or about 5 times the number of chemical synapses. Cp's were slightly larger in Drosophila than in Musca, although the Musca retinular axon has twice the diameter and length of that of the fruit fly. The evidence was reviewed in light of the likely supportive function of capitate projections on the R1-6 terminals.