Influence of spatial frequency on tuning and bias for orientation and direction in the cat's striate cortex.

Directionality, orientation and spatial frequency tuning were determined for 108 neurones recorded extracellularly from the striate cortex of anaesthetized cats. Significant sharpening of orientation selectivity with increasing spatial frequency was seen in all simple neurones and the overwhelming majority of complex neurones. Orientation selectivity sharpened in 90 and broadened in only 10 of 100 fully characterized neurones. At least four distinct classes of neurone could be characterized on the basis of their directionality at optimal spatial frequency, and the presence or absence of changes in directionality over a range of spatial frequencies: in two classes, directionality was spatial-frequency dependent; in the remaining two it was invariant. With two exceptions Type A neurones (23 cells) were direction-selective; they were narrowly tuned for orientation and spatial frequency, and their directionality was invariant with spatial-frequency. The majority of neurones (52 cells) were Type B, most of which were direction-biased; their bias for direction varied systematically with spatial frequency. Type C were direction-biased and spatial-frequency selective (5 cells), but showed a clear reversal of bias with change in spatial frequency. Type D, a subset of direction-biased cells, were bidirectional and spatial-frequency invariant (8 cells), with comparable response strengths to motion in two opposing directions at all spatial frequencies. These response types crossed traditional boundaries between categories of simple and complex neurones, assigned on the basis of spatial summation, presence or absence of end-inhibition, and receptive field size.