Relationship between orientation domains, cytochrome oxidase stripes, and intrinsic horizontal connections in squirrel monkey area V2.

Area V2, the main target of primary visual cortex projections, is characterized by a striking functional and connectional compartmentalization. Many aspects of this organization are correlated to three sets of stripes (thick, thin, and pale) revealed by cytochrome oxidase (CO) staining. Several questions related to the physiological properties of these compartments, their intrinsic connections, and points of similarity with area V1 modules are still unresolved. We have addressed some of these questions by combining the techniques of optical imaging of intrinsic signals, tract tracing, and CO histochemistry in the same patches of areas V1 and V2 of the squirrel monkey. The following observations were made. Orientation domains: in area V1 these are organized in narrow bands, while in area V2 they form patches. In area V2, domain width and distance between domains are approximately double that found in area V1. Orientation and CO stripe organization: orientation tuning was organized so that highly selective regions were centered on thick CO stripes while regions of broad orientation selectivity were centered on thin CO stripes. However, the orientation domains appeared to ignore borders between thick and pale stripes. Intrinsic connections: injections of the sensitive tracer biocytin into area V2 labeled a dense network of horizontally projecting fibers that were organized in columnar patches. Patches were small (mean width, 211 microns; mean length, 342 microns) and the labeling pattern extended over 4-5 mm. Axonal patches and CO stripes: Axonal patches found were in all three stripe compartments. However, injections that straddled the borders of thick/pale stripe compartments produced axonal projections that tended to cluster around border regions. Axonal patches and orientation domains: V2 injections produced labeling in V1 that appeared to be organized in narrow bands, reminiscent of orientation domain distribution in V1. Within area V2, axonal patches targeted a wide range of orientation domains, but appeared to avoid domains having orthogonal orientation preference to that found at the injection site. To conclude, our results show, on the one hand, a measure of functional specificity for the CO stripes and the intrinsic connections. On the other hand, they indicate additional substructures within area V2, whose precise relationship to the known compartmental organization remains to be clarified.