Representation of the brain's superior protocerebrum of the flesh fly, Neobellieria bullata, in the central body.

The central complex of the insect brain is a system of midline neuropils involved in transforming sensory information into behavioral outputs. Genetic studies focusing on nerve cells supplying the central complex from the protocerebrum propose that such neurons play key roles in circuits involved in learning the distinction of visual cues during operant conditioning. To better identify the possible sites of such circuits we used Bodian and anti-synapsin staining to resolve divisions of the superior protocerebrum into discrete neuropils. Here we show that in the fly Neobellieria bullata, the superior protocerebrum is composed of at least five clearly defined regions that correspond to those identified in Drosophila melanogaster. Intracellular dye fills and Golgi impregnations resolve "tangential neurons" that have intricate systems of branches in two of these regions. The branches are elaborate, decorated with specializations indicative of pre- and postsynaptic sites. The tangentially arranged terminals of these neurons extend across characteristic levels of the central complex's fan-shaped body. In this and another blowfly species, we identify an asymmetric pair of neuropils situated deep in the fan-shaped body, called the asymmetric bodies because of their likely homology with similar elements in Drosophila. One of the pair of bodies receives collaterals from symmetric arrangements of tangential neuron terminals. Cobalt injections reveal that the superior protocerebrum is richly supplied with local interneurons that are likely participants in microcircuitry associated with the distal processes of tangential neurons. Understanding the morphologies and arrangements of these and other neurons is essential for correctly interpreting functional attributes of the central complex.