Specificity in the Development of Vertical Connections in Cat Striate Cortex

The main efferent axons of pyramidal cells in layer 2/3 in the adult cat striate cortex make collateral connections specifically within layer 2/3 and layer 5 and avoid the intervening layer 4. Intracellular dye injections in vitro were used to determine how, during early postnatal development, this precise pattern of laminar connections was achieved. These investigations revealed that the pattern of collateral outgrowth was specific from the very earliest time that axons began sprouting collaterals. During the first postnatal week, sprouts were seen exclusively within layers 2/3 and 5; no evidence for a transient connection to layer 4 was observed. Furthermore, collaterals emerged simultaneously within layers 2/3 and 5, despite the large difference in the postmigratory ages of the two layers. By the end of the second postnatal week, the adult number of collaterals was achieved. Further elaboration of the local arbors occurred by repeated branching of already existing collaterals, rather than by addition of new collaterals to the main axon. These results demonstrate that the formation of local connections between cortical layers is highly specific, in contrast to the development of clustered horizontal connections by these same cells within layers 2/3 and 5, which involves extensive remodelling of local connections.