Local circuit neurons of macaque monkey striate cortex: III. Neurons of laminae 4B, 4A, and 3B.

We continue an investigation of the organization of local circuit neurons (largely inhibitory, GABAergic neurons, with smooth or sparsely spined dendrites) in the primary visual cortex of macaque monkey (Lund, '87: J. Comp. Neurol. 257:60-92; Lund et al., '88: J. Comp. Neurol. 276:1-29). This account covers local circuit neurons of layers 4B, 4A, and 3B; these three layers each receive different intrinsic second-order relays of principal thalamic inputs as well as receiving primary thalamic inputs in the case of two of the three laminae (4A and 3B). The study shows the existence of a number of different local circuit neurons making interlaminar projections between 4B, 4A, and 3B; each provides specific cross links between different combinations of the three laminae. It is known that the functional properties recorded physiologically from layers 4B, 4A, and 3B differ from one another and so these anatomical cross links may allow for correlation between different attributes of visual stimuli, e.g., color or motion, while still enabling separate processing of these different attributes to proceed in each of the three layers and be passed on to extrastriate areas. Whereas no spine-bearing neurons of layers 4B, 4A, or 3B provide "feedback" circuits to layer 4C (the source of their major intrinsic excitatory afferents), some of the local circuit neurons provide precisely structured axon feedback projections to divisions of 4C. The local circuit neurons also project to either lamina 5 or lamina 6, but not both and to superficial layers 3A, 2, and 1. Some local circuit neuron axon projections are of a dimension that would be confined to single functional clusters, e.g., cytochrome-rich "blobs," others reach out far enough to contact nearest neighbor "unlike" functional clusters, and yet others spread far enough to link repeating clusters of single function.