From neural arbors to daisies

Pyramidal neurons in layers 2 and 3 of the neocortex collectively form an horizontal lattice of long-range, periodic axonal projections, known as the superficial patch system. The precise pattern of projections varies between cortical areas, but the patch system has nevertheless been observed in every area of cortex in which it has been sought, in many higher mammals. Although the clustered axonal arbors of single pyramidal cells have been examined in detail, the precise rules by which these neurons collectively merge their arbors remain unknown. To discover these rules, we generated models of clustered axonal arbors following simple geometric patterns. We found that models assuming spatially aligned but independent formation of each axonal arbor do not produce patchy labeling patterns for large simulated injections into populations of generated axonal arbors. In contrast, a model that used information distributed across the cortical sheet to generate axonal projections reproduced every observed quality of cortical labeling patterns. We conclude that the patch system cannot be built during development using only information intrinsic to single neurons. Information shared across the population of patch-projecting neurons is required for the patch system to reach its adult state.