An anatomical substrate for the inhibitory gradient in the VLVp of the owl

The interaural difference in the level of sounds is an important cue for the localization of the sound's source. In the barn owl, a keen auditory predator, this binaural cue is first computed in the nucleus ventralis lemnisci laterale, pars posterior (VLVp), a cell group found within the fibers of the lateral lemniscus. Its neurons are excited by inputs from the contralateral ear and inhibited by inputs to the ipsilateral ear and are therefore sensitive indicators of interaural level difference. The excitation arrives by a direct input from the contralateral nucleus angularis, a cochlear nucleus, and the inhibition is mediated by a commissural projection that interconnects the VLVps of the two sides. The dorsally located neurons in the VLVp are more heavily inhibited than those found more ventrally, thus giving rise to a gradient of inhibition. This inhibitory gradient plays a central role in recent models of VLVp function. We present evidence based on standard anterograde tracing methods that this gradient of inhibition is mediated by a dorsoventral gradient in the density of synaptic inputs from the contralateral VLVp, the source of inhibition. Specifically, injection of tracers into one VLVp, regardless of the position of the injection within the nucleus, produced a vertically oriented field of label that was densest along the dorsal margin of the contralateral VLVp and became sparser a more ventral levels. Furthermore, we found that injections into the medial and lateral aspects of the nucleus produced this dorsoventrally graded field of label along the medial and lateral aspects of the contralateral VLVp, respectively. Finally, we confirmed an earlier observation suggesting that the anterior and posterior aspects of one VLVp project to the anterior and posterior aspects of the contralateral nucleus, respectively.