Prominent excitatory pathways in the cat visual cortex (A 17 and A 18): A current source density analysis of electrically evoked potentials

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Abstract
The current source density (CSD) method in its one-dimensional approximation is used to analyze the field potentials in visual areas 18 and 17 of the cat, which were elicited by stimulating electrodes in the optic chiasm (OX), the optic radiation (OR) or in the respective cortical area itself. The CSD analysis reveals the basic pattern of excitatory postsynaptic activity. In both visual areas the basic specific excitatory activity flows along three different intracortical pathways, all starting in layer IV: The first pathway relays activity from layer IV to supragranular pyramidal cells via strong, local connections to layer III and from there through long-distance connections to layer II. The second pathway conveys activity from layer IV to layer V, where it mainly contacts apical dendrites of layer VI pyramidal cells. This infragranular polysynaptic activity is not clearly resolvable into separate components, suggesting that it is conveyed by various groups of axons, among them long-distance horizontal connections. The third pathway has one synaptic relay within layer IV and then conveys activity to layer III. In addition, monosynaptic activity is revealed in layers VI and I. In A 18 one coherent, fast-conducting group of afferents induces this basic activity pattern. In A 17 no such fast conducting input is resolvable; the supragranular activity is induced by a small group of afferents with intermediate conduction velocity, which terminate in the upper part of layer IV. The infragranular activity is induced by afferents with slower and widely scattered conduction velocities, which terminate in the lower part of layer IV. The layer VI input is very prominent in A 17 and also has a wide latency scatter. The supragranular activity is more prominent in A 18 than in A 17 and the respective layers appear thicker, in accordance with anatomy. In A 17 the infragranular activity prevails and layers IV and VI appear very broad, again in accordance with anatomy. Comparison of the CSDs with the original evoked potentials shows that the surface evoked potentials over A 18 reflect the three dipolar sink/source distributions of the coherent monosynaptic activity in layer IV and of the two prominent polysynaptic activities in layers III and II. The widely scattered activity in the lower part of layer IV in A 17 and all infragranular activities in both areas generate smaller, partly closed-field potentials; those are not discernible from the strong far-field potentials which originate from the supragranular activity and — especially in A 17 —from farther distant events.