Laminar organization of working memory signals in human visual cortex

The human primary visual cortex (V1) is not only activated by incoming visual information, but is also engaged by top-down cognitive processes, such as visual working memory, even in the absence of visual input. This feedback is likely critical to our ability to visualize specific visual features, as higher-order cognitive control regions lack the visual selectivity to represent such information. Clearly, such internally generated signals do not trigger genuine perception of the remembered stimulus, meaning they must be organized in a manner that is different to bottom-up driven signals. Internally generated signals may be kept separate from incoming sensory data by virtue of the laminar organization of inter-area cortical connections. Namely, bottom-up driving connections target layer 4, located in the middle of the cortical column, while feedback connections target deep and superficial layers and largely avoid layer 4. Using lamina-resolved fMRI, we simultaneously measured the activity in three early visual cortical areas (V1-V3) that are recruited to represent stimulus information during visual working memory. We observed item-specific working memory signals in early visual cortex. In V1, this item-specific activity was selectively present at deep and superficial cortical depths, avoiding the middle layers, while working memory related activity was present at all depths in V2 and V3. These results show for the first time the laminar organization of internally generated signals during visual working memory in the human visual system, and provide new insights into how bottom-up and top-down signals in visual cortex are deployed.