Morphological basis for nonsynaptic communication within the central nervous system by exocytotic release of secretory material from the egg-laying stimulating neuroendocrine caudodorsal cells of Lymnaea stagnalis.

The fine structure of the axons of the cerebral, egg-laying stimulating caudodorsal cells of the snail Lymnaea stagnalis has been studied with various light and electron microscope techniques. Special attention was paid to exocytotic release of secretory material (demonstrated with the tanic acid method) from nonsynaptic release sites in the cerebral commissure. This phenomenon has been compared with neurohaemal release. The commissure consists of two morphological compartments, separated by a sheath of glial cells. The outer compartment is formed by the neurohaemal area of the caudodorsal cells, the inner consists of thousands of, mainly unidentified, axons. Furthermore, ventral caudodorsal cells send axons through the inner compartment. These give rise to collaterals, which divide into smaller collaterals, forming an extensive network ("collateral system") throughout the inner compartment. Eventually, collaterals end blindly within the inner compartment. They contain the same three morphological types of secretory granule as the neurohaemal axon terminals. The collaterals never form synaptic contacts; exocytotic release of the contents of secretory granules takes place at nonsynaptic release sites. These sites occur rather dispersed and do not face one particular type of neighbouring neural element. As in the neurohaemal area, both single and multiple exocytoses occur. Widened intercellular spaces, filled with flocculent, electron-dense material, occur near highly active nonsynaptic release sites. The spaces are often bordered by glial cells and may facilitate diffusion of released secretory material through the inner compartment. Apparently, a ventral caudodorsal cell releases secretory material in two fashions: from neurohaemal axon terminals into the haemolymph, and nonsynaptically, from the collaterals into the intercellular space of the central nervous system. Possible functions of the glial sheath between the neurohaemal area and the inner compartment are proposed. Most likely, the collateral system enables the caudodorsal cells to communicate with targets within the central nervous system in a nonsynaptic fashion. A possible target is the cerebral Ring Neuron, which sends an axon branch through the inner compartment and, as was previously shown neurophysiologically, is controlled by the caudodorsal cells in a nonsynaptic fashion.