Two sets of wide-field neurons extend neurites into the fly's optic lamina, where monopolar cells receive photoreceptor input. They exhibit immunoreactivity to antibodies raised against either 5-hydroxytryptamine or the crustacean peptide PDH, respectively. Both are proposed whole-field neuromodulators of vision, apparently regulating a circadian rhythm of monopolar cell size. Seeking functional correlates, we have re-examined the electroretinogram for circadian rhythmicity, and for responses to locally injected 5-hydroxytryptamine and peptide. Long-term electroretinogram recordings from Calliphora entrained to a light/dark cycle and then transferred to constant darkness, uncovered a gradual, modest increase during the subjective night in the electroretinogram's ON- and OFF-transients, from the lamina's monopolar cells. Five to twenty nl of 5-hydroxytryptamine (10(-3) mol.1(-1)) injected into the head haemolymph strongly enhanced the electroretinogram transients, an action reversed by 5-hydroxytryptamine antagonists. Injected into the eye, 5-hydroxytryptamine (10(-4) mol.1(-1)) had the opposite effect; the rapid onset there suggests direct action, whilst the opposing effect from haemolymph injection suggests a different receptor site. Pigment-dispersing hormone (2.2 x 10(-5) mol.1(-1)) injected into the haemolymph increased the electroretinogram transients along a biphasic course, with a slow partial recovery; injected into the eye, it lacked effect.

Musca and related flies have three main photoreceptor subsystems. The R1-6 group has short axons that terminate in the cartridges of the first optic neuropile, the lamina. The cartridges are bypassed by the longer axons of R7 and R8, which run together to terminate at different levels in the underlying medulla neuropile. The present account describes a shallow, previously unidentified zone in the lamina within which R7/8 make glancing contact with R1-6. At the distal border of the cartridge over no more than 3-4 microns depth, the tangentially directed short axon of R6 squeezes between the pair from R7 and R8, forming quite large areas of mutual contact (approximately 7 microns2). Less frequently, R1 is contacted. At least some of these sites contain smaller membrane specialisations indistinguishable from the more numerous gap junctions found more proximally that interconnect the terminals of R1-6. The R7/8 junctions with R6 are of comparable size (0.15 micron 2) and likewise possess symmetrical membrane densities. They provide proposed pathways for direct electrical interaction to account for observed electrical input from R7/8 to the R1-6 subsystem. In two cases R7/8 was possibly postsynaptic to R1-6 at a multiple-contact synapse, but even if functional, these sites were so rare that they are unlikely to have much operational significance.

No abstract

A long-term objective of our studies on the first optic neuropil (or lamina) underlying the fly's compound eye is to explore how afferent photoreceptor synapses disappear during normal adult experience. To increase the frequency of this loss and the chances for its detection artificially, we have examined in this study the synapses during the degeneration of their presynaptic elements, the synaptic terminals of the receptor cells. This may be reliably procured by illuminating for 12 min with strong green light eyes that have received an injection of the dye sulforhodamine 101 (Picaud et al., 1988). The lesion is local and develops rapidly. Degeneration among terminals is progressive but asynchronous. There are several different types of degeneration, most interpretable as stages in a temporal progression after illumination-induced injury. Degenerative changes include shrinkage and darkening of terminals and mitochondrial swelling. Synaptic sites are lost in a defined sequence: (1) the T-shaped presynaptic ribbon disappears first; (2) the members of what is normally a tetrad of postsynaptic elements withdraw as an ensemble from the receptor terminal's membrane, and the surrounding epithelial glial cells extend between former pre- and postsynaptic partners; and (3) the postsynaptic elements then separate from each other. In the most rapidly affected terminals, the frequencies for those synaptic sites at which both presynaptic ribbons and postsynaptic elements remain intact decline by 85%, even in the first 8 hr postillumination.

The cellular mechanisms by which nervous systems evolve to match evolutionary changes occurring in the rest of the body remain largely unexplored. In a distal visual neuropil of a previously unexamined ancient dipteran family, Stratiomyidae, homologues of all of the periodic neurons known already from more recent Diptera can be recognized, occupying the same locations within the unit structure. This points to extreme developmental stasis for more than 200 million years, conserving both cell identity and position. The arborizations that some neurons make also have remained conservative, but others show marked differences between families in both size and branching patterns. At the electron-microscopical level, extensive differences in synaptic connectivity are found, some sufficient to radically redefine the systems roles of particular neurons. The findings bear out an earlier prediction that changes in the connectivity matrix linking conserved neurons may have been a major factor in implementing evolutionary change in the nervous system.