Golgi and EM studies of the formation of dendritic and axonal arbors: The interneurons of the substantia gelatinosa of rolando in newborn kittens

Golgi studies in newborn kittens show that the two most prevalent interneurons in Rexed's lamina II of the dorsal horn of the medulla, the stalked cell and islet cell (Gobel, 1975a,b, 1978b) form their dendritic arbors in a similar fashion. At birth, both cell types are present in forms ranging from immature, in which numerous short dendrites radiate from the cell body in all directions, to relatively mature in which their dendritic arbors have elongated in specific directions and the adult branching pattern is already evident. During postnatal maturation, many dendrites are lost while only a few go on to lengthen. The unmyelinated axons of both cells are first recognized in forms in which lengthening dendrites have taken on their preferred direction of orientation. The two parts of Rexed's lamina II, i.e., layers IIa and IIb have already reached their adult mediolateral width at birth and the neuropil has nearly achieved its adult compactness. Space in the compact neuropil for elongating neuronal and astrocytic processes becomes available through the disintegration of many existing dendrites and by an overall fourfold increase in the rostrocaudal length of the dorsal horn of the medulla during postnatal maturation. At birth, the lengthening of the plasma membranes of elongating neuronal and astrocytic processes proceeds as vesicles (addition vesicles) found in aggregates throughout dendrites, unmyelinated axons and astrocytic processes fuse with and become incorporated into the existing plasma membranes. In addition, many dendrites in layers IIa and IIb are beading up and disintegrating. Within the beads, neurotubules are lost and addition vesicles fuse with each other to form small cavities. These cavities continue to enlarge, hollowing out the beads. The cavities ultimately open to the intercellular space as their membranes fuse with the plasma membrane of the beads. Finally, the beads disintegrate and their plasma membranes fragment. The thread‐like segments between adjacent disintegrating beads shrivel until they ultimately disappear. Disintegration of beaded dendrites results in very little debris and does not provoke a phagocytic glial reaction. The disintegration of a dendritic branch takes place in spite of synaptic input from non‐primary axons and is thought to occur from a failure to establish synaptic connections with primary axonal endings.