Multiple visual areas in the caudal superior temporal sulcus of the macaque

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Abstract
Anatomical and physiological evidence indicates that, in addition to area MT, much of the cortex in the caudal superior temporal sulcus (STS) of the macaque has visual functions. Yet the organization of areas outside of MT remains unclear, and there are even conflicting data on the boundaries of MT itself. To examine these issues, we recorded from neurons throughout this region in three monkeys. Anterograde or retrograde tracers were injected into MT at the conclusion of recording to identify its projection fields. Based on differences in their visuotopic organization, neuronal properties, receptive field size, myeloarchitecture, and pattern of connections with MT, several visual areas were distinguished within the caudal STS.Area MT, defined as the heavily myelinated portion of the striate (VI) projection zone in STS, contained a systematic representation of only about the central 30°–40° of the contralateral field. The far peripheral field was represented medial to MT in MTp, which we had previously found receives projections from far peripheral V1 and V2 (Ungerleider and Desimone: J. Comp. Neurol 248:147–163, 1986). Like MT, MTp contained a high proportion of directionally selective cells, and receptive field size in MTp was the size expected of MT fields if the latter were to extend into the periphery.Areas MST (medial superior temporal) and PP (posterior parietal) were found medial to MT and MTp. Both MST and PP had a high proportion of directionally selective cells, but only MST received a direct projection from MT. Cells in MST had larger receptive fields than those in either MT or MTp butnonetheless displayed a crude visuotopic organization. Receptive fields of cells in PP were even larger, some including the entire contralateral visual field. Furthermore, unlike cells in MST, some in PP responded to auditoryor somesthetic stimuli in addition to visual stimuli.Area FST, which has a distinctive myeloarchitecture, was found anterior to MT in the fundus of the STS, for which it is named. FST received a direct projection from MT, but only about a third of its cells were directionally selective. Receptive fields of cells in FST were large, often included the center of gaze, and often crossed into the ipsilateral visual field.Area V4t (transitional V4) and a portion of V4 were found lateral to MT within the STS, and both received direct projections from MT. V4t has a distinctive, light myelination. Both areas had a low incidence of directionally selective cells, and both contained coarse representations of the lower visual field.The neuronal properties of areas in the caudal STS suggest that MT, MTp, MST, and PP, together with the superior temporal polysensory area, constitute a cortical system for motion analysis. At successive stages in this system, neurons appear to integrate motion information over an increasingly large portion of the retina, respond selectively to more complex types of motion, and respond to inputs from additional sensory modalities.