Translaminar circuits formed by the pyramidal cells in the superficial layers of cat visual cortex.

Pyramidal cells in the superficial layers of the neocortex provide a major excitatory projection to layer 5, which contains the pyramidal cells that project to subcortical motor-related targets. Both structurally and functionally rather little is known about this interlaminar pathway, especially in higher mammals. Here, we made sparse ultrastructural reconstructions of the projection to layer 5 of three pyramidal neurons from layer 3 in cat V1 whose morphology, physiology, and synaptic connections with layers 2 and 3 were known. The dominant targets of the 74 identified synapses in layer 5 were the dendritic spines of pyramidal cells. The fractions of target spiny dendrites were 59, 61, and 84% for the three cells, with the remaining targets being dendrites of smooth neurons. These fractions were similar to the distribution of targets of unlabeled asymmetric synapses in the surrounding neuropil. Serial section reconstructions revealed that the target dendrites were heterogenous in morphology, indicating that different cell types are innervated. This new evidence indicates that the descending projection from the superficial layer pyramidal cells does not simply drive the output pyramidal cells that project to cortical and subcortical targets, but participates in the complex circuitry of the deep cortical layers.

Synaptic connections formed by patchy projections of pyramidal cells in the superficial layers of cat visual cortex.

The present study is the first to describe quantitatively the patterns of synaptic connections made by the patchy network of pyramidal cell axons in the superficial layers of cat V1 in relation to the orientation map. Intrinsic signal imaging of the orientation map was combined with 3D morphological reconstructions of physiologically-characterized neurons at light and electron microscope levels. A Similarity Index (SI) expressed the similarity of the orientation domain of a given bouton cluster to that of its parent dendritic tree. Six pyramidal cells whose axons had a wide range of SIs were examined. Boutons were sampled from five local and five distal clusters, and from the linear segments that link the clusters. The synaptic targets were reconstructed by serial section electron microscopy. Of the 233 synapses examined, 182 synapses were formed with spiny neurons, the remainder with smooth neurons. The proportion of smooth neurons that were synaptic targets varied greatly (from 0 to 50%) between the cluster samples, but was not correlated with the SI. The postsynaptic density sizes were similar for synapses in local and distal clusters, regardless of their SI. This heterogeneity in the synaptic targets of single cells within the superficial layers is a network feature well-suited for context-dependent processing.