The connections of the middle temporal visual area (MT) and their relationship to a cortical hierarchy in the macaque monkey

The cortical and subcortical connections of the middle temporal visual area (MT) of the macaque monkey were investigated using combined injections of 3Hproline and horseradish peroxidase within MT. Cortical connections were assigned to specific visual areas on the basis of their relationship to the pattern of interhemispheric connections, revealed by staining for degeneration following callosal transection. MT was shown to be reciprocally connected with many topographically organized cortical visual areas, including V1, V2, V3, and V4. These pathways link regions representing corresponding portions of the visual field in the different areas. In addition, MT has reciprocal connections with two previously unidentified cortical areas, which we have designated the medial superior temporal area (MST) and the ventral intraparietal area (VIP). The laminar distribution of terminals and cell bodies in cortical areas connected with MT follows a consistent pattern. In areas V1, V2, and V3, the projections to MT arise largely or exclusively from cells in supragranular layers, and the reciprocal connections from MT terminate mainly in supragranular and infragranular layers. In contrast, the projections to MST and VIP terminate mainly in layer IV, and the reciprocal pathways originate from cells in both superficial and deep layers. On the basis of this pattern, each connection can be designated as forward or feedback in nature, and a hierarchical arrangement of visual areas can be determined. In this hierarchy, MT is at a higher level than V1, V2, and V3, and at a lower level than MST and VIP. Subcortical projections were seen from MT to the claustrum, the putamen, the caudate nucleus, the inferior and lateral subdivisions of the pulvinar complex, the ventral lateral geniculate nucleus, the reticular nucleus of the thalamus, the superior colliculus, and the pontine nuclei.