The working memory networks of the human brain.

Working memory and short-term memory are closely related in their cognitive architecture, capacity limitations, and functional neuroanatomy, which only partly overlap with those of long-term memory. The author reviews the functional neuroimaging literature on the commonalities and differences between working memory and short-term memory and the interplay of areas with modality-specific and supramodal representations in the brain networks supporting these fundamental cognitive processes. Sensory stores in the visual, auditory, and somatosensory cortex play a role in short-term memory, but supramodal parietal and frontal areas are often recruited as well. Classical working memory operations such as manipulation, protection against interference, or updating almost certainly require at least some degree of prefrontal support, but many pure maintenance tasks involve these areas as well. Although it seems that activity shifts from more posterior regions during encoding to more anterior regions during delay, some studies reported sustained delay activity in sensory areas as well. This spatiotemporal complexity of the short-term memory/working memory networks is mirrored in the activation patterns that may explain capacity constraints, which, although most prominent in the parietal cortex, seem to be pervasive across sensory and premotor areas. Finally, the author highlights open questions for cognitive neuroscience research of working memory, such as that of the mechanisms for integrating different types of content (binding) or those providing the link to long-term memory.