Morphological types of projection neurons in layer 5 of cat visual cortex.

Pyramidal cells in layer 5 of the visual cortex have multiple cortical and subcortical projection sites. Previous studies found that many cells possess bifurcating axons and innervate more than one cortical or subcortical target, but cells projecting to both cortical and subcortical targets were not observed. The present study examines the morphology of cells in cat visual cortex projecting to the superior colliculus, the main subcortical target of layer 5, and cells in layer 5 projecting to cortical areas 18 and 19. The neurons that give rise to these different projections were retrogradely labelled and intracellularly stained in living brain slices. Our results show that cells within each projection group have several morphological features in common. All corticotectal cells have a long apical dendrite forming a large terminal tuft in layer 1. Their cell bodies are medium sized to large, and their basal dendrites form a dense and symmetrical dendritic field. Corticocortical cells in layer 5 have a very different morphology: their apical dendrites are short and they never reach higher than layers 2/3. Their cells bodies are small to medium sized and they have fewer basal dendrites than corticotectal cells. Thus there are two morphologically distinct projection systems in layer 5, one projecting to cortical and the other one to subcortical targets, suggesting that these two systems transmit different information from the visual cortex. Among the corticotectal cells with the largest cell bodies we found some cells whose basal and apical dendrites were almost devoid of spines. Spiny and spinefree corticotectal cells also have different intrinsic axon collaterals and therefore play different roles in the cortical circuitry. While many spiny corticotectal cells have axon collaterals that project to layer 6, spinefree corticotectal cells have fewer axon collaterals and these do not arborize in layer 6. We suggest that the two morphological types of corticotectal cells might be related to functional differences known to exist among these cells. We discuss how the presence or absence of spines affects the integration of the synaptic input and how this might be related to the cells' functional properties.