Development of axonal arbors of layer 6 pyramidal neurons in ferret primary visual cortex.

We followed the development of axonal arbors of layer 6 pyramidal neurons in ferret striate cortex to determine whether early developing axon collaterals are formed specifically in the correct target layers from the outset or achieve their adult specificity by the elimination of initially exuberant projections. These neurons were chosen for study because they are amongst the first to be generated in the developing ferret's striate cortex, and, in mature animals, these cells have axonal arbors that are highly specific for layer 4 and to a lesser extent layers 2/3 but have few collateral branches in layer 5. The axonal arbors of individual layer 6 pyramidal neurons were reconstructed following labeling in living slices prepared from the striate cortex of ferrets aged 13-35 days postnatal (P13-35). At the earliest ages (P13-15), axonal arbors consisted of a simple axon extending from the base of the cell body into the subplate or white matter and usually forming a few collateral branches but never ascending into layer 5. By P19-20, about one-half of the cells had extended axon collaterals into layer 5 or higher, and these already appeared to branch preferentially in layer 4. All of the cells from older animals had substantial axonal arbors in layers 2-4. By P26-28, there were approximately ten times as many axonal branches in layer 4 as in layer 5. Between P26-28 and P35, there was no significant change in the number of branches in layer 5, but the numbers of both branches and of axon collateral terminations in layer 4 approximately doubled. Thus, the extent of axonal arborization in layer 4 increases dramatically between P13 and P35, and growth is highly specific for correct target layers, with few branches formed in layer 5.