Long-range horizontal connections between supragranular pyramidal cells in the extrastriate visual cortex of the rat

In this study, we examined the morphological structure and synaptic physiology of long-range axon projections among supragranular pyramidal cells in the extrastriate visual cortex of the rat. Intra- and extracellular recordings from layer II/III pyramidal cells were performed in brain slices of area 18a following extracellular stimulation of either the underlying white matter or within layer II/III. Neurons were injected with biocytin for two-dimensional reconstruction of their axon arborizations. The conduction velocity of afferent fibers (0.58 m/s) was twice as high as that of intracortical tangential fibers (0.28 m/s). Layer II/III cells were mainly di- or polysynaptically driven by afferent activation, but predominantly monosynaptically driven from intracortical stimulation sites. The afferent as well as intracortically evoked postsynaptic potentials showed a very similar time course and shape. From both stimulation sites, suprathreshold action potentials could be elicited. The current threshold for a postsynaptic response and the slope and width of excitatory postsynaptic potentials (EPSPs) increased with the distance of lateral stimulation. The morphological properties of layer II/III pyramidal cell axon collaterals closely corresponded to the electrophysiological results. Long-range intraareal axon collaterals could be followed up to 1 mm within the supragranular layers. Their length-distance distribution showed an inverse relationship to the threshold currents of EPSPs. Pyramidal cells exhibited regularly spaced patches of horizontal axon collaterals with an interpatch distance of about 250 microns. We concluded that the supragranular horizontal network in the extrastriate visual cortex of the rat is qualitatively very similar to that of cats and monkeys. However, quantitative differences exist in its spatial extent and physiological characteristics.