Effects of spatial attention on contrast response functions in macaque area V4

Previous single-unit studies of visual cortex have reported that spatial attention modulates responses to different orientations and directions proportionally, such that it does not change the width of tuning functions for these properties. Other studies have suggested that spatial attention causes a leftward shift in contrast response functions, such that its effects on responses to stimuli of different contrasts are not proportional. We have further explored the effects of attention on stimulus-response functions by measuring the responses of 131 individual V4 neurons in two monkeys while they did a task that controlled their spatial attention. Each neuron was tested with a set of stimuli that spanned complete ranges of orientation and contrast during different states of attention. Consistent with earlier reports, attention scaled responses to preferred and nonpreferred orientations proportionally. However, we did not find compelling evidence that the effects were best described by a leftward shift of the contrast response function. The modulation of neuronal responses by attention was well described by either a leftward shift or proportional scaling of the contrast response function. Consideration of differences in experimental design and analysis that may have contributed to this discrepancy suggests that it was premature to exclude a proportional scaling of responses to different contrasts by attention in favor of a leftward shift of contrast response functions. The current results reopen the possibility that the effects of attention on stimulus-response functions are well described by a single proportional increase in a neuron's response to all stimuli.