Reorganization of Exuberant Axonal Arbors Contributes to the Development of Laminar Specificity in Ferret Visual Cortex

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Abstract
Layer-specific cortical axons are believed to develop precisely from the outset without making exuberant branches in incorrect cortical layers. We tested this hypothesis by following the development of axonal arbors of layer 2/3 pyramidal neurons in ferret visual cortex, which in the adult have dense axonal arbors in layers 2/3 and 5 and not in layer 4. We began our studies at earlier ages and sampled at closer intervals than in previous studies. Our studies reveal that between postnatal day (P) 14 and P18 the initial growth of axonal arbors lacks specificity for layers 2/3 and 5 and involves the formation of a small number of branches incorrectly in layer 4. After P18 the number of incorrect axonal branches in layer 4 decreased, whereas there was extensive axonal growth specifically in layers 2/3 and 5. To further study the mechanisms involved in directing the layer-specific growth and elimination of axonal branches, we studied the development of layer 2/3 neurons in slices kept in culture for 5 d. In these studies the initial imprecise growth at P14–18 followed by precise growth after P18 was mimicked in vitro, but the elimination of incorrect axons from layer 4 did not occur. These studies reveal that most axonal arbors grow precisely in the correct layers, but the initial growth involves the formation of a small number of incorrect axonal branches that are later eliminated. Therefore the mechanisms that regulate axonal growth are not as precise initially as later, and mechanisms must exist to allow selective elimination of axon collaterals from incorrect layers.