Quantitative studies of single-cell properties in monkey striate cortex. I. Spatiotemporal organization of receptive fields.

1. The properties of single cells in striate cortex of the rhesus monkey, representing the visual field 2 degrees -5 degrees from the fovea, were examined quantitatively with stationary and moving stimuli. Three distinct classes of cells were identified: S type, CX type, and T type. 2. S-type cells were defined as those oriented cells which to the optimal direction of movement in their receptive fields exhibited one or more spatially separate subfields within each of which a response was obtained to either a light or dark edge, but not to both. Several different types of S-cells were distinguished: a) S1-type cells for which moving edges revealed a single excitatory area within which a response was elicited by either a light or a dark edge but not by both. Most of these cells were unidirectional. b) S2-type cells for which moving edges revealed two spatially separate response areas, one of which was excited by a light edge and the other by a dark edge. Both regions responded to the same direction of movement. c) S3-type cells which had two response areas, one of which was excited by a stimulus moving in one direction (at right angles to the axis of orientation) and the other, of opposite contrast, which responded in the opposite direction, d) S4-type cells which to one direction of movement showed two spatially separate regions sensitive to a light and dark edge and which in the other direction of movement had only one responsive area (either light or dark). e) Cells which had multiple spatially separate subfields (S5-7 types). 3. CX-type cells were defined as those oriented cells which in their receptive fields exhibited no spatial separation for light- and dark-edge responses; they discharged to both edges in the same direction of movement and in the same spatial area. Flashing stimuli elicited both on and off responses throughout the receptive field. CX-type cells were predominantly of two types: those which were selective for direction of stimulus movement and those which were not. 4. A third class of cells (T-type) were those which were excited by only one sign of contrast change and responded in a sustained fashion even when there was no contour within the receptive field. These cells were poorly or not at all oriented; some of them were selective to wavelength. 5. Quantitative comparisons showed the following differences between S-type and CX-type cells: a) S-type cells had smaller receptive fields than CX-type cells but the populations over-lapped considerably. Receptive-field size was smallest in layer 4c. In all other layers S-type cells had the same size fields. CX-type cells, by contrast, tended to have larger fields in layer 5-6 than 2-3. b) The spatial separation between light and dark response areas was the best criterion for distinguishing S-type and CX-type cells. The distribution of this measure disclosed two populations of cells with relatively limited overlap. c) In layers 2 and 3, both S-type and CX-type cells had low spontaneous activity...