The organization of double bouquet cells in monkey striate cortex

In previous publications we proposed a model of cortical organization in which the pyramidal cells of the cerebral cortex are organized into modules. The modules are centred around the clusters of apical dendrites that originate from the layer 5 pyramidal cells. In monkey striate cortex such modules have an average diameter of 23 μm and the outputs originating from the modules are contained in the vertical bundles of myelinated axons that traverse the deeper layers of the cortex. The present study is concerned with how the double bouquet cells in layer 2/3 of striate cortex relate to these pyramidal cell modules. The double bouquet cells are visualized with an antibody to calbindin, and it has been shown that their vertically oriented axons, or horse tails, are arranged in a regular array, such that there is one horse tail per pyramidal cell module. Within layer 2/3 the double bouquet cell axons run alongside the apical dendritic clusters, while in layer 4C they are closely associated with the myelinated axon bundles. However, the apical dendrites are not the principal targets of the double bouquet cell axons. Most of the neuronal elements post-synaptic to them are the shafts of small dendrites (60%) and dendritic spines, with which they form symmetric synapses. This regular arrangement of the axons of the double-bouquet cells and their relationship to the components of the pyramidal cells modules supports the concept that there are basic, repeating neuronal circuits in the cortex.