Synaptic biology of barrel cortex circuit assembly

Mature neuronal circuits arise from the coordinated interplay of cell-intrinsic differentiation programs, target-derived signals and activity-dependent processes. Typically, cell-intrinsic mechanisms predominate at early stages of differentiation, while input-dependent processes modulate circuit formation at later stages of development. The whisker barrel cortex of rodents is particularly well suited to study this latter phase. During the first few days after birth, thalamocortical axons (TCA) from the somatosensory ventral posteromedial nucleus (VPM) form synapses onto layer 4 (L4) neurons, which aggregate to form barrels, whose spatial organization corresponds to the distribution of the whiskers on the snout. Besides specific genetic programs, which control TCA and L4 neuron specification, the establishment of the barrel pattern also depends on the information resulting from whisker activation. The plasticity of this system during the first few days after birth is critical for barrel formation: damage to the sensory periphery impairs TCA patterning, while lesions after this period have less pronounced effects. Here, we will review the role and position of L4 neurons within cortical columnar circuits and synaptogenesis during barrel formation.