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The depth profiles of visually evoked field potentials were recorded in areas 17 and 18 of the cat visual cortex. For comparison, potential profiles evoked by electrical stimulation of the primary afferents and of the nonspecific reticular system were also recorded. From these profiles the current source-density (CSD) distributions were calculated using the one-dimensional CSD method. CSD distributions evoked by the different types of stimuli differ in their amplitudes and time courses by approximately one and two orders of magnitude. Qualitatively, however, they are very similar. Thus, the CSDs can be interpreted as reflecting the same basic pattern of excitatory synaptic activations. This pattern consists of early activation components in the input layers, followed by excitatory synaptic activations in layer III, then in layer II, and in layer V. The basic pattern of cortical activation was found to be modulated by specific features of the visual stimuli. Modulations reflecting contour-versus contrast-contents as well as those reflecting characteristic features of moving patterns have been identified. Most of the CSD components of the cortical activation sequence were obtained from regions extending well beyond the cellular receptive fields in visual cortex. Thus, they reflect nonretinotopic activities. Parameters other than specific features of the visual stimuli have profound influence on cortical CSDs. Nonspecific parameters which have been considered are the general state of cortical excitability, the temporal interactions of successive activities (which are predominantly facilitatory), and the lateral interactions of simultaneous activations from different regions of the visual field (predominantly inhibitory).

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. 1. 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. 2. 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. 3. 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. 4. 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.

The spatio-temporal distributions of excitatory synaptic ensemble activities in A17 and A18 of the visual cortex of the macaque monkey have been investigated. The synaptic activities were elicited by electrical stimulation of the primary efferents and were localized by applying the current source density analysis to the intracortically recorded field potentials. The principal results are as follows: 1. In A17, two groups of activity, evoked by fast and slow afferents, respectively, were distinguishable. 2. The fast afferents induced monosynaptic activity in layer IV C alpha and layer VI, disynaptic activity in layer IV C alpha and in the supragranular layers and trisynaptic activity in layer IV B. 3. The slow efferents induced monosynaptic activity in lower layers IV C beta and layer VI, disynaptic activity via strong connections in upper layer IV C beta, further disynaptic activity in layers III and IV B and trisynaptic activity in layers V A and II. 4. With the exception that the CSD data reveal more polysynaptic activity within layer IV, there is good agreement between the spatio-temporal distribution of synaptic activities and the cortical circuit diagrams proposed in anatomical studies. 5. In A18, activities from the slow and fast conducting afferent systems are revealed in layer IV, both most likely mediated by the monosynaptically activated target cells of A17. These activities are passed on to the supra-and infragranular layers. 6. In the lateral geniculate nucleus the safety factor of transmission is higher for activity conveyed by slow-than by fast-conducting retinal afferents. 7. The spatial distribution of monocularly evoked surface potentials failed to reveal the ocular dominance columns. 8. Comparison with the cat indicates that, with respect to the intracortical circuitry and LHN-transmission, there are more similarities between the fast-group activity in the monkey and the y-system in the cat and between the slow-group activity in the monkey and the x-system in the cat than vice versa.

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