Neocortical Layer 1: An Elegant Solution to Top-Down and Bottom-Up Integration

Annu Rev Neurosci. 2021 Jul 8;44:221-252. doi: 10.1146/annurev-neuro-100520-012117. Epub 2021 Mar 17.


Many of our daily activities, such as riding a bike to work or reading a book in a noisy cafe, and highly skilled activities, such as a professional playing a tennis match or a violin concerto, depend upon the ability of the brain to quickly make moment-to-moment adjustments to our behavior in response to the results of our actions. Particularly, they depend upon the ability of the neocortex to integrate the information provided by the sensory organs (bottom-up information) with internally generated signals such as expectations or attentional signals (top-down information). This integration occurs in pyramidal cells (PCs) and their long apical dendrite, which branches extensively into a dendritic tuft in layer 1 (L1). The outermost layer of the neocortex, L1 is highly conserved across cortical areas and species. Importantly, L1 is the predominant input layer for top-down information, relayed by a rich, dense mesh of long-range projections that provide signals to the tuft branches of the PCs. Here, we discuss recent progress in our understanding of the composition of L1 and review evidence that L1 processing contributes to functions such as sensory perception, cross-modal integration, controlling states of consciousness, attention, and learning.

Keywords: GABAergic interneurons; layer 1; neocortex; predictive coding; pyramidal cell dendrites; top-down processing.

Publication types

  • Research Support, N.I.H., Extramural

MeSH terms

  • Dendrites
  • Learning
  • Neocortex*
  • Pyramidal Cells