Membrane specializations in the first optic neuropil of the housefly, Musca domestica L. I. Junctions between neurons.

Abstract

Thin section and freeze-fracture replicas of the first optic neuropil (lamina ganglionaris) of the fly Musca were studied to determine the types, extent and location of membrane specializations between neurons. Five junctional types are found, exclusive of chemical synapses. These are gap, tight and septate junctions, close appositions between retinular (R) axons and capitate projections (in which an epithelial glial cell invaginates into an R axon). Junctional types and their cellular associations follow: gap junctions, between lamina (L) interneurons, L1-L2; tight junctions, between L1-L2; L3-L4; L4-epithelial glial cell; and R7-R8. Septate junctions, between L1-L2, L3-L4, L3-beta, L4-beta, alpha-beta, and an unidentified fibre making septate junctions with L1 and L2. Close appositions are found between R axons in the distal portion of the optic cartridges of this neuropil prior to extensive R chemical synapses with L1, L2. These loci (seen in freeze-fracture replicas) have rhomboidal patches of hexagonally arrayed P face particles. Intermembranous clefts between R axons are about 50 A and are invariably electron lucent. These points of near contact between R terminals are probably the sites of low electrical resistance measured by Shaw (1979). Capitate projections are for the first time revealed in freeze fracture surfaces. Here epithelial glia send many, short, mushroom-shaped processes invaginating into R axons forming a tenacious structural bond. All four membrane leaflets (P and E faces of R axon and glial membrane) in the capitate projection possess particles in higher densities than in the surrounding nonspecialized regions. The known, general functions of each membrane specialization were correlated with the functional capacities of those lamina neurons possessing them in an effort to interpret better the integrative capacity of this neuropil. These data provide some fine structural bases for a putative 'blood-brain' barrier between lamina and haemolymph, between lamina and peripheral retina, and possibly between lamina and second optic neuropil.