Visual system of calliphorid flies: organization of optic glomeruli and their lobula complex efferents.

Reconstructions of silver-stained brains revealed 27 optic glomeruli that occupy a major volume of the lateral protocerebrum. Axons from different morphological types of columnar output neurons from the lobula complex sort out to specific glomeruli. Glomeruli are partially enwrapped by glial processes and are invaded by the dendrites and terminals of local interneurons that connect different glomeruli in a manner analogous to local interneurons in the antennal lobes. Each type of columnar neuron contributes to a palisade-like ensemble that extends across the whole or a circumscribed area of the retinotopic mosaic. A second class of outputs from the lobula comprises wide-field neurons, the dendrites of which interact with planar fields or column-like patches of retinotopic inputs from the medulla. These neurons also send their axons to optic glomeruli. Dye fills demonstrate that lobula complex neurons supplying glomeruli do not generally terminate directly on descending neurons. Local interneurons and projection neurons provide integrative circuitry within and among glomeruli. As exemplified by the anterior optic tubercle, optic glomeruli can also have elaborate internal architectures. The results are discussed with respect to the identification of motion- and orientation-selective neurons at the level of the lobula and lateral protocerebrum and with respect to the evolutionary implications raised by the existence of neural arrangements serving the compound eyes, which are organized like neuropils serving segmental ganglia equipped with appendages.