Convergence of retinal W-cell and corticotectal input to cells of the cat superior colliculus

1. Conduction velocities of retinotectal W-cell afferents were estimated from differences among latencies of collicular unit responses to supramaximal stimulation of the contralateral optic disk (OD), optic chiasm (OX), and ipsilateral optic tract (OT). W-cell afferents driving collicular neurons had very slowly conducting axons, nearly all below 8 m/s (mean = 5.3 m/s). These match the conduction velocities of W-cell axons terminating in the uppermost superficial gray layer and triggering juxtazonal potentials (JZPs). Such slow conduction velocities are typical of W-cells belonging to the W2 subclass ("phasic W-cells"), but are slower than nearly all W1 cells ("tonic W-cells"). 2. Most W-driven cells were activated at latencies longer than expected for monosynaptic input from these W-cell afferents. However, comparable delays were observed among JZPs, which signal monosynaptic excitation of collicular neurons by W-cell terminals. This suggests that the delayed activation of W-driven cells reflects slowed conduction in the preterminal segments of W-cell afferents rather than polysynaptic transmission of W-cell signals through intermediary neurons in the brain stem or cortex. Thus monosynaptic inputs from retinal W2 cells appear to drive most neurons of the superficial collicular layers. 3. Convergence of retinotectal W-cell and corticotectal pathways was assessed by recording responses of W-driven collicular cells to intracortical stimulation of area 17. The great majority of W-driven collicular cells were activated by cortical stimulation (41/52; 79%), indicating that such convergence is widespread. 4. The population of corticotectal cells influencing W-driven collicular cells may differ from that mediating Hoffmann's Y-indirect pathway. W-driven collicular cells were activated from the striate cortex at longer latencies (mean = 6.3 ms) than cells driven by the Y-indirect pathway (mean = 2.5 ms). In addition, cortically activated W-driven cells were common throughout the superficial gray layer, whereas cells driven by the Y-indirect input were encountered only in the deepest part of the superficial gray and below. 5. W2 cells, apparently the dominant retinotectal cell type, nearly all project contralaterally and are tuned for slow stimulus velocities. Thus the binocularity of W-driven collicular cells and their sensitivity to moderately fast stimulus motion probably reflect the convergent cortical input described here.