Topography of attention in the primary visual cortex

Previous research suggests that feedback circuits mediate the effect of attention to the primary visual cortex (V1). This inference is mainly based on temporal information of the responses, where late modulation is associated with feedback signals. However, temporal data alone are inconclusive because the anatomical hierarchy between cortical areas differs significantly from the temporal sequence of activation. In the current work, we relied on recent physiological and computational models of V1 network architecture, which have shown that the thalamic feedforward, local horizontal and feedback contribution are reflected in the spatial spread of responses. We used multifocal functional localizer and quantitative analysis in functional magnetic resonance imaging to determine the spatial scales of attention and sensory responses. Representations of 60 visual field regions in V1 were functionally localized and four of these regions were targets in a subsequent attention experiment, where human volunteers fixated centrally and performed a visual discrimination task at the attended location. Attention enhanced the peak amplitudes significantly more in the lower than in the upper visual field. This enhancement by attention spread with a 2.4 times larger radius (approximately 10 mm, assuming an average magnification factor) compared with the unattended response. The corresponding target region of interest was on average 20% stronger than that caused by the afferent sensory stimulation alone. This modulation could not be attributed to eye movements. Given the contemporary view of primate V1 connections, the activation spread along the cortex provides further evidence that the signal enhancement by spatial attention is dependent on feedback circuits.