In the present work we propose a new phylogenetic hypothesis for the role played by cortical and subcortical afferents to the nucleus of the optical tract, the main visual relay station of the horizontal optokinetic reflex in mammals. The hypothesis is supported by anatomical and physiological data obtained in the South American opossum ( Didelphis aurita) using the following experimental approaches: (i) single-unit recordings in the nucleus of the optic tract and simultaneous electrical stimulation of the contralateral nucleus of the optic tract; (ii) single-unit recordings in the nucleus of the optic tract and simultaneous electrical stimulation of the ipsilateral striate cortex; (iii) injection of cholera toxin subunit B into the striate cortex and subsequent immunohistochemical reaction to reveal the presence of the marker in the thalamus and mesencephalon; and (iv) single-unit recordings in the nucleus of the optic tract both before and after ablation of the ipsilateral visual cortex. The main results are: (i) there is a strong inhibitory reciprocal effect upon the nucleus of the optic tract following stimulation of its contralateral counterpart; (ii) electrophysiological and anatomical data imply that the visual cortex does not project directly to the nucleus of the optic tract. Rather, cortical terminals seem to target the nearby anterior and posterior pretectal nuclei and orthodromic latencies in the nucleus of the optic tract following stimulation of the visual cortex were twice as large as in the superior colicullus; and (iii) ablation of the entire visual cortex did not have any effect upon binocularity of cells in the nucleus of the optic tract. These results strengthen the model proposed here for the role of the interactions between the nuclei of the optic tract under optokinetic stimulation. The hypothesis in the present work is that the cortical influences upon the nucleus of the optical tract, in addition to the subcortical ones, appeared only recently in phylogenesis. In more primitive mammals, such as the opossum, subcortical interactions are thought to play a relatively important role. With the emergence of retinal specializations, such as the fovea, one might suppose that there followed the appearance of new ocular movements, such as the smooth pursuit and certain types of saccades, that came to join the pre-existent optokinetic reflex.
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