SummaryA fundamental challenge for the visual system is to integrate visual features into a coherent scene, known as the binding problem. The neural mechanisms of feature binding are hard to identify because of difficulties in separating active feature binding from feature co-occurrence. In previous studies on feature binding [1–5], visual features were superimposed and presented simultaneously. Neurons throughout the visual cortex are known to code multiple features [6]. Therefore, the observed binding effects could be due to the physical co-occurrence of features and the sensory representation of feature pairings. It is uncertain whether the mechanisms responsible for perceptual binding were actually recruited [7, 8]. To address this issu...