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The lateral superior olive (LSO) is believed to encode differences in sound level at the two ears, a cue for azimuthal sound location. Most high-frequency-sensitive LSO neurons are binaural, receiving inputs from both ears. An inhibitory input from the contralateral ear, via the medial nucleus of the trapezoid body (MNTB), and excitatory input from the ipsilateral ear enable level differences to be encoded. However, the classical descriptions of low-frequency-sensitive neurons report primarily monaural cells with no contralateral inhibition. Anatomical and physiological evidence, however, shows that low-frequency LSO neurons receive low-frequency inhibitory input from ipsilateral MNTB, which in turn receives excitatory input from the contralateral cochlear nucleus and low-frequency excitatory input from the ipsilateral cochlear nucleus. Therefore, these neurons would be expected to be binaural with contralateral inhibition. Here, we re-examined binaural interaction in low-frequency (less than ∼3 kHz) LSO neurons and phase locking in the MNTB. Phase locking to low-frequency tones in MNTB and ipsilaterally driven LSO neurons with frequency sensitivities <1.2 kHz was enhanced relative to the auditory nerve. Moreover, most low-frequency LSO neurons exhibited contralateral inhibition: ipsilaterally driven responses were suppressed by raising the level of the contralateral stimulus; most neurons were sensitive to interaural time delays in pure tone and noise stimuli such that inhibition was nearly maximal when the stimuli were presented to the ears in-phase. The data demonstrate that low-frequency LSO neurons of cat are not monaural and can exhibit contralateral inhibition like their high-frequency counterparts. [ABSTRACT FROM AUTHOR]

Recent psychoacoustic experiments have shown that interaural time differences provide adequate cues for lateralizing high-frequency sounds, provided the stimuli are complex and not pure tones. We present here physiological evidence in support of these findings. Neurons of high best frequency in the cat inferior colliculus respond to interaural phase differences of amplitude modulated waveforms, and this response depends upon preservation of phase information of the modulating signal. Interaural phase differences were introduced in two ways: by interaural delays of the entire waveform and by binaural beats in which there was an interaural frequency difference in the modulating waveform. Results obtained with these two methods are similar. Our results show that high-frequency cells can respond to interaural time differences of amplitude modulated signals and that they do so by a sensitivity to interaural phase differences of the modulating waveform. [ABSTRACT FROM AUTHOR]

Spatial hearing, and more specifically the ability to localize sounds in space, is one of the most studied and best understood aspects of hearing. Because there is no coding of acoustic space at the receptor organ, physiological sensitivity to spatial aspects of sounds first emerges in the central nervous system. Much progress has been made in the identification and characterization of the circuits in the auditory brainstem that create sensitivity to binaural and monaural cues toward acoustic space. We review the progress over the past third of a century, with a focus on the mammalian brainstem and on the anatomy and cellular physiology underlying the physiological tuning of monaural and binaural circuits to acoustic cues toward spatial hearing. In addition to examining the detailed mechanisms involved in the processing of the three main spatial cues, we also review the integration of these cues and their use toward behavior. © 2019 American Physiological Society. Compr Physiol 9:1503-1575, 2019., (Copyright © 2019 American Physiological Society. All rights reserved.)

Almost all behavioral studies of sound localization have used either an approach-to-target or pointing/orienting task to assess absolute sound localization performance, yet there are very few direct comparisons of these measures. In an approach-to-target task, the subject is trained to walk to a sound source from a fixed location. In an orienting task, finger, head and/or eye movements are monitored while the subject's body is typically constrained. The fact that subjects may also initiate head and eye movements toward the target during the approach-to-target task allows us to measure the accuracy of the initial orienting response and compare it with subsequent target selection. To perform this comparison, we trained cats to localize a broadband noise presented randomly from one of four speakers located ± 30° and ± 60° in azimuth. The cat responded to each sound presentation by walking to and pressing a lever at the perceived location, and a food reward was delivered if the first attempt was correct. In tandem, we recorded initial head and eye orienting movements, via magnetic search coils, immediately following target onset and prior to the walking response. Reducing either stimulus duration or level resulted in a systematic decline in both measurements of localization performance. When the task was easy, localization performance was accurate for both measures. When the task was more difficult, the number of incorrect (i.e., wrong selection) and no-go (i.e., no selection) responses increased. Interestingly, for many of the incorrect trials, there was a dissociation between the orienting response and the target selected, and for many of the no-go trials, the gaze oriented towards the correct target even though the cat did not move to it. This suggests different neural systems governing walking to a target as compared to unconditioned gaze orienting., (Copyright © 2014 Elsevier B.V. All rights reserved.)

Studies of sound localization use relative or absolute psychoacoustic paradigms. Relative tasks assess acuity by determining the smallest angle separating two sources that subjects can discriminate, the minimum audible angle (MAA), whereas absolute tasks measure subjects' abilities to indicate sound location. It is unclear whether or how measures from the two tasks are related, though the belief that the MAA is specifically related to the precision of absolute localization is common. The present study aimed to investigate the basis of this relationship by comparing the precision of absolute location estimates with a measure of spatial acuity computed from the same data. Three cats were trained to indicate apparent sound source locations that varied in azimuth and elevation via orienting gaze shifts (combined eye and head movements). The precision of these absolute responses, as measured by their standard deviation, was compared with acuity thresholds derived from receiver operating characteristic (ROC) analyses of the cumulative distributions. Surprisingly, the acuity measures were occasionally very poor indicators of absolute localization precision. Incongruent results were attributed to errors in mean accuracy, which are disregarded in analyses of traditional relative tasks. Discussion focuses on the potential for internal biases to affect measures of localization acuity.

As an animal navigates its surroundings, the sounds reaching its two ears change in waveform similarity (interaural correlation) and in time of arrival (interaural time difference, ITD). Humans are exquisitely sensitive to these binaural cues, and it is generally agreed that this sensitivity involves coincidence detectors and internal delays that compensate for external acoustic delays (ITDs). Recent data show an unexpected relationship between the tuning of a neuron to frequency and to ITD, leading to several proposals for sources of internal delay and the neural coding of interaural temporal cues. We review the alternatives, and argue that an understanding of binaural mechanisms requires consideration of sensitivity not only to ITDs, but also to interaural correlation.

The precedence effect (PE) describes several spatial perceptual phenomena that occur when similar sounds are presented from two different locations and separated by a delay. The mechanisms that produce the effect are thought to be responsible for the ability to localize sounds in reverberant environments. Although the physiological bases for the PE have been studied, little is known about how these sounds are localized by species other than humans. Here we used the search coil technique to measure the eye positions of cats trained to saccade to the apparent locations of sounds. To study the PE, brief broadband stimuli were presented from two locations, with a delay between their onsets; the delayed sound meant to simulate a single reflection. Although the cats accurately localized single sources, the apparent locations of the paired sources depended on the delay. First, the cats exhibited summing localization, the perception of a "phantom" sound located between the sources, for delays < +/-400 micros for sources positioned in azimuth along the horizontal plane, but not for sources positioned in elevation along the sagittal plane. Second, consistent with localization dominance, for delays from 400 micros to about 10 ms, the cats oriented toward the leading source location only, with little influence of the lagging source, both for horizontally and vertically placed sources. Finally, the echo threshold was reached for delays >10 ms, where the cats first began to orient to the lagging source on some trials. These data reveal that cats experience the PE phenomena similarly to humans.

Mammals localize sound sources in azimuth based on two binaural cues, interaural differences in the time of arrival and level of the sounds at the ears. In contrast, the cue for elevation is based on patterns of the broadband power spectra at each ear that result from the direction-dependent acoustic filtering properties of the head and pinnae. Although the exact form of this "spectral shape" cue is unknown, most attention has been directed toward a prominent direction-dependent energy minimum, or "notch," because its location in frequency, for both humans and cats, moves predictably from low to high as a source is moved from low to high elevations. However, there is little direct evidence that these spectral notches are important elevational cues in animals other than humans. Here we demonstrate a striking illusion in the localization of sounds in elevation by cats using stimulus configurations that elicit summing localization and the precedence effect that can be explained by spectral shape cues.