Synaptic remodelling during development and maturation: junction differentiation and splitting as a mechanism for modifying connectivity.

Morphological variation of the synaptic active zone during later development and maturation (15-224 days) has been studied in the molecular layer of the rat occipital cortex. Both E-PTA stained and osmicated tissue have been used. In the E-PTA stained material the degree of specialization of the presynaptic thickening is directly related to junction length. Junctions with well-developed dense projections (Types A and B) are longest and continue to increase in length with maturation, suggesting that active remodelling of the synaptic apposition is an ongoing process. The presence of perforated junctions, possessing two or more regions with specializations of different maturity and different curvature, raises the possibility that these junctions may arise by the addition and differentiation of new paramembranous material at an existing junction. In osmicated tissue, presynaptic terminals possessing multiple active zones have been quantitated. The maturational increase in number of simple perforated junctions (Type 1), is paralleled by a smaller increase in the number of multiple perforated junctions (Type 2). A spine apparatus is frequently observed in these perforated terminals, suggesting that it is intimately involved in the reorganization. The direction of curvature of the closely apposed junctions is predominantly negative (indenting the postsynaptic process). Other types of arrangement, with separate postsynaptic processes, are described (Types 3-5), and micrographs suggestive of sequential stages in pre- and postsynaptic terminal splitting are presented. The total length of the postsynaptic thickening of perforated terminals is twice the mean synaptic length of non-perforated terminals, again suggestive that duplication of the active zone may have occurred. Division of existing synaptic terminals by duplication and subsequent splitting would readily account for the increased dendritic spinal numbers seen in Golgi preparations of animals raised under enriched conditions. This would be a straight-forward mechanism by which reinforcement of neuronal connections could occur in response to use.