Organization of the zona incerta in the macaque: a Nissl and Golgi study

The zona incerta has been implicated in the control of the initiation of saccadic eye movements in the primate. Complex interactions within the zona incerta must take place to integrate its varied inputs and to produce a coherent efferent signal in order for this function to occur. However, whether the anatomical substrates exist within the zona incerta to allow this integration to take place has not been established. The zona incerta in monkeys (Macaca mulatta) was examined in frontally, horizontally, and sagittally sectioned preparations stained for Nissl, myelinated fibers, or cytochrome oxidase, or impregnated by the Golgi technique. This nucleus can be separated into dorsal and ventral laminae on the basis of staining and morphological differences between these two subdivisions. Neurons are more densely packed, more darkly stained, and larger in the ventral lamina. In addition, the neuropil of the ventral lamina is much more intensely stained after cytochrome oxidase histochemistry. Two neuronal types, principal cells and interneurons, were identified on the basis of neuronal cell body, dendritic, and axonal features in Golgi-impregnated preparations. Principal cells have fusiform or polygonal somata (long axis from 18 to 40 microns) and dendrites that extend for up to 750 microns within the lamina in which the cell bodies are located. Putative local interneurons have small (12-16 microns), round or oval cell bodies with wavy dendrites (up to 400 microns). Numerous multilobed appendages and axon-like processes originate from these dendrites and make apparent contacts with other interneurons or with dendrites of principal cells. Dendrites of most neurons in both laminae are oriented preferentially along the principal axis, dorsolateral-to-ventromedial, of the nucleus. Therefore, within the limits of light microscopy, the zona incerta appears to possess the morphological heterogeneity to form complex intrinsic interactions. These interactions are hypothesized to form the integrative substrate for the large array of incertal inputs that are utilized to produce an efferent signal involved in the initiation of saccadic eye movements.