Orientation and Direction Selectivity of Neurons in V1 of Alert Monkeys: Functional Relationships and Laminar Distributions

We studied orientation selectivity in V1 of alert monkeys and its relationship to other physiological parameters and to anatomical organization. Single neurons were stimulated with drifting bars or with sinusoidal gratings while compensating for eye position. Orientation selectivity based on spike counts was quantified by circular variance and by the bandwidth of the orientation tuning curve. The circular variance distribution was bimodal, suggesting groups with low and with high selectivity. Orientation selectivity was clearly correlated with spontaneous activity, classical receptive field (CRF) size and the strength of surround suppression. Laminar distributions of neuronal properties were distinct. Neurons in the output layers 2/3, 4B and 5 had low spontaneous activity, small CRFs and high orientation selectivity, while the input layers had greater diversity. Direction-selective cells were among the neurons most selective for orientation and most had small CRFs. A narrow band of direction- and orientation-selective cells with small CRFs was located in the middle of layer 4C, indicating appearance of very selective cells at an early stage of cortical processing. We suggest that these results reflect interactions between excitatory and inhibitory mechanisms specific to each sublamina. Regions with less inhibition have higher spontaneous activity, larger CRFs and broader orientation tuning. Where inhibition is stronger, spontaneous activity almost disappears, CRFs shrink, and orientation selectivity is high.