Physiology and Impact of Horizontal Connections in Rat Neocortex

Cortical information processing at the cellular level has predominantly been studied in local networks, which are dominated by strong vertical connectivity between layers. However, recent studies suggest that the bulkof axonstargeting pyramidal neurons most likelyoriginatefromoutsidethislocal range,emphasizingtheimportance of horizontal connections.We mappeda subset of these connections to L5B pyramidal neurons in rat somatosensory cortex with photostimulation, identifying intact projections up to a lateral distance of 2 mm. Our estimates of the spatial distribution of cells presynaptic to L5B pyramids support the idea that the majority is located outside the local volume. The synaptic physiology of horizontal connections does not differ markedly from that of local connections, whereas the layer and cell-type-dependent pattern of innervation does. Apart from L2/3, L6A provides a strong source of horizontal connections. Implementing our data into a spiking neuronal networkmodelshowsthatmore horizontal connectionspromoterobust asynchronousongoing activitystates and reducenoise correlations in stimulus-induced activity.