Covert enaction at work: Recording the continuous movements of visuospatial attention to visible or imagined targets by means of Steady-State Visual Evoked Potentials (SSVEPs)

Abstract

Whereas overt visuospatial attention is customarily measured with eye tracking, covert attention is assessed by various methods. Here we exploited Steady-State Visual Evoked Potentials (SSVEPs) - the oscillatory responses of the visual cortex to incoming flickering stimuli - to record the movements of covert visuospatial attention in a way operatively similar to eye tracking (attention tracking), which allowed us to compare motion observation and motion extrapolation with and without eye movements. Observers fixated a central dot and covertly tracked a target oscillating horizontally and sinusoidally. In the background, the left and the right halves of the screen flickered at two different frequencies, generating two SSVEPs in occipital regions whose size varied reciprocally as observers attended to the moving target. The two signals were combined into a single quantity that was modulated at the target frequency in a quasi-sinusoidal way, often clearly visible in single trials. The modulation continued almost unchanged when the target was switched off and observers mentally extrapolated its motion in imagery, and also when observers pointed their finger at the moving target during covert tracking, or imagined doing so. The amplitude of modulation during covert tracking was ∼25-30% of that measured when observers followed the target with their eyes. We used 4 electrodes in parieto-occipital areas, but similar results were achieved with a single electrode in Oz. In a second experiment we tested ramp and step motion. During overt tracking, SSVEPs were remarkably accurate, showing both saccadic-like and smooth pursuit-like modulations of cortical responsiveness, although during covert tracking the modulation deteriorated. Covert tracking was better with sinusoidal motion than ramp motion, and better with moving targets than stationary ones. The clear modulation of cortical responsiveness recorded during both overt and covert tracking, identical for motion observation and motion extrapolation, suggests to include covert attention movements in enactive theories of mental imagery.

Keywords: Mental imagery; Motion extrapolation; SSVEPs; Smooth pursuit eye movements; Visuospatial attention.