Interaural time difference is an important cue for sound localization. In the barn owl (Tyto alba) neuronal sensitivity to this disparity originates in the brainstem nucleus laminaris, Afferents from the ipsilateral and contralateral magnocellular cochlear nuclei enter the nucleus laminaris through its dorsal and ventral surfaces, respectively, and interdigitate in the nucleus. Intracellular recordings from these afferents show orderly changes in conduction delay with depth in the nucleus. These changes are comparable to the range of interaural time differences available to the owl. Thus, these afferent axons act as delay lines and provide anatomical and physiological bases for a neuronal map of interaural time differences in the nucleus laminaris.Jeffress (1) proposed a model, the place theory, for measurement of the interaural time differences that underlie sound localization. His theory contained two important concepts, the first being the principles of delay lines and coincidence detection. The second states that the "place" or anatomical location of the neuron in an array encodes the place or location of the sound. In the model, the coincidence detectors are binaural neurons which require simultaneous arrival of spikes from the two sides to elicit a maximal discharge. Signals reach these coincidence detectors by axonal paths which are unequal for the two sides. This difference in path lengths translates into a disparity in spike conduction time. The coincidence detectors respond preferentially to an interaural time difference that delays the arrival of spikes on the shorter path, and advances the arrival of spikes on the longer path so as to cause the spikes to arrive simultaneously. Subsequently, several authors proposed cross-correlation models of sound localization using the same principles (2, 3). Goldberg and Brown (4), working on the superior olive ofdogs, obtained data consistent with the Jeffress model, and similar results have been obtained in cats (5).Barn owls use interaural time differences to localize sound in azimuth (6). Neuronal sensitivity to these time differences arises in the nucleus laminaris (7), the avian homologue of the medial superior olive. As in the superior olive, physiological responses from nucleus laminaris are consistent with the principles of the Jeffress model (8).Despite the widespread acceptance of the Jeffress model, the nature of the delay lines and the cellular mechanisms of coincidence detection remain to be investigated. Conduction delays can be produced by several different methods, such as variation in membrane time constants, number of synapses, length of dendrites, and length of axonal path (9, 10). We present both anatomical and physiological evidence for axonal delay lines in the barn owl's nucleus laminaris.MATERIALS AND METHODS Anatomy. Horseradish peroxidase (HRP) was used as a tracer to study the innervation of nucleus laminaris in five owls (Tyto alba) (11). Detailed protocols may be found in ref.12. Birds were anesthetized by intramuscular (i.m.)...