Attentional modulation of visual responses by flexible input gain.

Although it is clear that sensory responses in the cortex can be strongly modulated by stimuli outside of classical receptive fields as well as by extraretinal signals such as attention and anticipation, the exact rules governing the neuronal integration of sensory and behavioral signals remain unclear. For example, most experiments studying sensory interactions have not explored attention, while most studies of attention have relied on the responses to relatively limited sets of stimuli. However, a recent study of V4 responses, in which location, orientation, and spatial attention were systematically varied, suggests that attention can both facilitate and suppress specific sensory inputs to a neuron according to behavioral relevance. To explore the implications of such input gain, we modeled the effects of a center-surround organization of attentional modulation using existing receptive field models of sensory integration. The model is consistent with behavioral measurements of a suppressive effect that surrounds the facilitatory locus of spatial attention. When this center-surround modulation is incorporated into realistic models of sensory integration, it is able to explain seemingly disparate observations of attentional effects in the neurophysiological literature, including spatial shifts in receptive field position and the preferential modulation of low contrast stimuli. The model is also consistent with recent formulations of attention to features in which gain is variably applied among cells with different receptive field properties. Consistent with functional imaging results, the model predicts that spatial attention effects will vary between different visual areas and suggests that attention may act through a common mechanism of selective and flexible gain throughout the visual system.