How 'visual' is the visual cortex? The interactions between the visual cortex and other sensory, motivational and motor systems as enabling factors for visual perception

The definition of the visual cortex is primarily based on the evidence that lesions of this area impair visual perception. However, this does not exclude that the visual cortex may process more information than of retinal origin alone, or that other brain structures contribute to vision. Indeed, research across the past decades has shown that non-visual information, such as neural activity related to reward expectation and value, locomotion, working memory and other sensory modalities, can modulate primary visual cortical responses to retinal inputs. Nevertheless, the function of this non-visual information is poorly understood. Here we review recent evidence, coming primarily from studies in rodents, arguing that non-visual and motor effects in visual cortex play a role in visual processing itself, for instance disentangling direct auditory effects on visual cortex from effects of sound-evoked orofacial movement. These findings are placed in a broader framework casting vision in terms of predictive processing under control of frontal, reward- and motor-related systems. In contrast to the prevalent notion that vision is exclusively constructed by the visual cortical system, we propose that visual percepts are generated by a larger network-the extended visual system-spanning other sensory cortices, supramodal areas and frontal systems.