STRUCTURAL ORGANIZATION OF THE SUPERIOR COLLICULUS

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Many types of experiments indicate that the superior colliculus plays a key role in initiating orienting movements of the head, eyes and ears toward objects of interest. At least three classes of models of the relationship between its structure and function can be distinguished: those proposing that pathways between the layers of the superior colliculus integrate the sensory and motor systems involved in initiating orienting movements; those arguing that the layers are independent and serve mainly to distribute information to different destinations; and those proposing that connections between compartments within individual layers provide the substrate for interactions between the sensory and motor systems. The specific aims of this application are to test these models. Because of technical limitations, past anatomical studies have provided few clues to intracollicular circuitry. Consequently, current models have been based primarily on physiological properties of collicular neurons, such as the latencies of their responses and properties of their receptive and movement fields. These models postulate specific patterns of connections, but the postulates are difficult to test with physiological methods. The proposed research will test the current models directly by intracellularly injecting cells of each layer and tracing their intracollicular connections, using living brain slices from the tree shrew, Tupaia belangeri. Relative to other commonly studied species, this primate-like mammal has an especially large and well differentiated superior colliculus, which greatly facilitates the task of analyzing the connections between the layers. In some cases, cells will be prelabeled by retrograde axonal transport so that the injected cells can be identified by the extracollicular destinations of their axons, in addition to their location, morphology and intracollicular connections. In other experiments, sensory or motor pathways to the superior colliculus will be prelabeled to determine whether they converge on collicular neurons of different types. These experiments are designed to provide a framework for new models based on knowledge of intracollicular circuitry and, in this way, will contribute to our understanding of the neural mechanisms underlying sensorimotor integration in the vertebrate brain.