Interaural time difference

Abstract: THE LOCALIZATION of high-frequency sinusoids depends on differences in the intensity of the sounds arriving at the two ears, that of low-frequency sinusoids on differences in phase (30). The present paper considers how the introduction of these two cues affects the discharge of individual binaural neurons of the superior olivary complex. Also described is the response of these cells to variations in the average intensity of binaural stimuli. Most of the neurons were located in the medial superior olive (MSO) or the medial preolivary nucleus (MPO).

Abstract: Detection of interaural time differences underlies azimuthal sound localization in the barn owl Tyto alba. Axons of the cochlear nucleus magnocellularis, and their targets in the binaural nucleus laminaris, form the circuit responsible for encoding these interaural time differences. The nucleus laminaris receives bilateral inputs from the cochlear nucleus magnocellularis such that axons from the ipsilateral cochlear nucleus enter the nucleus laminaris dorsally, while contralateral axons enter from the ventral side. This interdigitating projection to the nucleus laminaris is tonotopic, and the afferents are both sharply tuned and matched in frequency to the neighboring afferents. Recordings of phase-locked spikes in the afferents show an orderly change in the arrival time of the spikes as a function of distance from the point of their entry into the nucleus laminaris. The same range of conduction time (160 mu sec) was found over the 700-mu m depth of the nucleus laminaris for all frequencies examined (4-7.5 kHz) and corresponds to the range of interaural time differences available to the barn owl. The estimated conduction velocity in the axons is low (3-5 m/sec) and may be regulated by short internodal distances (60 mu m) within the nucleus laminaris. Neurons of the nucleus laminaris have large somata and very short dendrites. These cells are frequency selective and phase-lock to both monaural and binaural stimuli. The arrival time of phase-locked spikes in many of these neurons differs between the ipsilateral and contralateral inputs. When this disparity is nullified by imposition of an appropriate interaural time difference, the neurons respond maximally. The number of spikes elicited in response to a favorable interaural time difference is roughly double that elicited by a monaural stimulus. Spike counts for unfavorable interaural time differences fall well below monaural response levels. These findings indicate that the magnocellular afferents work as delay lines, and the laminaris neurons work as co- incidence detectors. The orderly distribution of conduction times, the predictability of favorable interaural time differences from monaural phase responses, and the pattern of the anatomical projection from the nucleus laminaris to the central nucleus of the inferior colliculus suggest that interaural time differences and their phase equivalents are mapped in each frequency band along the dorsoventral axis of the nucleus laminaris.

Abstract: Two experiments are described in which listeners judge the apparent directions of virtual sound sources–headphone‐presented sounds that are processed in order to simulate free‐field sounds. Previous results suggest that when the cues to sound direction are preserved by the simulation, the apparent directions of virtual sources are nearly the same as the apparent directions of real free‐field sources. In the experiments reported here, the interaural phase relations in the processing algorithms are manipulated in order to produce stimuli in which the interaural time difference cues signal one direction and interaural intensity and pinna cues signal another direction. The apparent directions of these conflicting cue stimuli almost always follow the interaural time cue, as long as the wideband stimuli include low frequencies. With low frequencies removed from the stimuli, the dominance of interaural time difference disappears, and apparent direction is determined primarily by interaural intensity difference and pinna cues.

Abstract: Microsecond differences in the arrival time of a sound at the two ears (interaural time differences, ITDs) are the main cue for localizing low-frequency sounds in space. Traditionally, ITDs are thought to be encoded by an array of coincidence-detector neurons, receiving excitatory inputs from the two ears via axons of variable length ('delay lines'), to create a topographic map of azimuthal auditory space(1,2). Compelling evidence for the existence of such a map in the mammalian ITD detector, the medial superior olive (MSO), however, is lacking. Equally puzzling is the role of a-temporally very precise(3)-glycine-mediated inhibitory input to MSO neurons. Using in vivo recordings from the MSO of the Mongolian gerbil, we found the responses of ITD-sensitive neurons to be inconsistent with the idea of a topographic map of auditory space. Moreover, local application of glycine and its antagonist strychnine by iontophoresis (through glass pipette electrodes, by means of an electric current) revealed that precisely timed glycine-controlled inhibition is a critical part of the mechanism by which the physiologically relevant range of ITDs is encoded in the MSO. A computer model, simulating the response of a coincidence-detector neuron with bilateral excitatory inputs and a temporally precise contralateral inhibitory input, supports this conclusion.