Spatial and temporal frequency selectivity of neurones in visual cortical areas V1 and V2 of the macaque monkey.

The spatial and temporal frequency selectivity of 148 neurones in the striate cortex, V1, and of 122 neurones in the second visual cortical area, V2, of the macaque monkey were studied using sine-wave gratings of suprathreshold contrast drifting over the receptive field at the preferred orientation and direction. Neurones in V1 and V2 were selective for different but partially overlapping ranges of the spatial frequency spectrum. At retinal eccentricities of 2-5 deg from the fovea, the spatial frequency preferences for neurones ranged from 0.5 to 8.0 cycles/deg in V1 and from 0.2 to 2.1 cycles/deg in V2 and were on average almost 2 octaves lower in V2 than in V1. Spatial frequency full band widths in the two cortical areas were in the range 0.8-3.0 octaves, with a mean value of 1.8 octaves, in the parafoveal representation of both V1 and V2, and 1.4 and 1.6 octaves respectively in the foveal representation of V1 and V2. Most neurones in V1 and some in V2 responded well at temporal frequencies up to 5.6-8.0 Hz before their responses dropped off at still higher frequencies. In V1, 68% of the neurones exhibited low-pass temporal tuning characteristics and 32% were very broadly tuned, with a mean temporal frequency full band width of 2.9 octaves. However, in V2 only 30% of the neurones showed low-pass temporal selectivity and 70% of the cells had bandpass temporal characteristics, with a mean full band width of 2.1 octaves. In V2 the minimal overlap of bandpass tuning curves across the temporal frequency spectrum suggests that there are at least two distinct bandpass temporal frequency mechanisms as well as neurones with low-pass temporal frequency tuning at each spatial frequency. A matrix of spatial and temporal frequency combinations was employed as stimuli for neurones with bandpass temporal frequency selectivity in both V1 and V2. The resultant spatio-temporal surfaces provided evidence that a neurone's preference for spatial frequency is essentially independent of the test temporal frequency; however, in V2 there was some tendency for temporal frequency peaks to shift slightly towards lower frequencies when non-optimum values of spatial frequency either above or below the preferred value were tested. Neurones with pronounced directional selectivity were encountered over a wide range of spatial frequencies, although in both cortical areas there was a tendency for an increased incidence of directional selectivity among neurones which were selective for lower spatial frequencies and higher temporal frequencies.