Neural Correlates of the Precedence Effect in the Inferior Colliculus of Behaving Cats

2004 ◽  
Vol 92 (6) ◽  
pp. 3286-3297 ◽  
Author(s):  
Daniel J. Tollin ◽  
Luis C. Populin ◽  
Tom C. T. Yin
Keyword(s):  
Inferior Colliculus ◽  
Sound Localization ◽  
Species Differences ◽  
Behavioral Responses ◽  
Neural Correlates ◽  
Precedence Effect ◽  
Single Neurons ◽  
Localization Dominance ◽  
Echo Threshold ◽  
Spatial Locations

Several auditory spatial illusions, collectively called the precedence effect (PE), occur when transient sounds are presented from two different spatial locations but separated in time by an interstimulus delay (ISD). For ISDs in the range of localization dominance (<10 ms), a single fused sound is typically located near the leading source location only, as if the location of the lagging source were suppressed. For longer ISDs, both the leading and lagging sources can be heard and localized, and the shortest ISD where this occurs is called the echo threshold. Previous physiological studies of the extracellular responses of single neurons in the inferior colliculus (IC) of anesthetized cats and unanesthetized rabbits with sounds known to elicit the PE have shown correlates of these phenomena though there were differences in the physiologically measured echo thresholds. Here we recorded in the IC of awake, behaving cats using stimuli that we have shown to evoke behavioral responses that are consistent with the precedence effect. For small ISDs, responses to the lag were reduced or eliminated consistent with psychophysical data showing that sound localization is based on the leading source. At longer ISDs, the responses to the lagging source recovered at ISDs comparable to psychophysically measured echo thresholds. Thus it appears that anesthesia, and not species differences, accounts for the discrepancies in the earlier studies.

scholarly journals Influence of Sound Source Location on the Behavior and Physiology of the Precedence Effect in Cats

2009 ◽  
Vol 102 (2) ◽  
pp. 724-734 ◽  
Author(s):  
Micheal L. Dent ◽  
Daniel J. Tollin ◽  
Tom C. T. Yin
Keyword(s):  
Auditory Nerve ◽  
Single Unit ◽  
Neural Correlates ◽  
Precedence Effect ◽  
Neural Responses ◽  
Localization Dominance ◽  
Single Unit Recordings ◽  
Echo Threshold ◽  
Unanesthetized Animals ◽  
Psychophysical Task

Psychophysical experiments on the precedence effect (PE) in cats have shown that they localize pairs of auditory stimuli presented from different locations in space based on the spatial position of the stimuli and the interstimulus delay (ISD) between the stimuli in a manner similar to humans. Cats exhibit localization dominance for pairs of transient stimuli with |ISDs| from ∼0.4 to 10 ms, summing localization for |ISDs| < 0.4 ms and breakdown of fusion for |ISDs| > 10 ms, which is the approximate echo threshold. The neural correlates to the PE have been described in both anesthetized and unanesthetized animals at many levels from auditory nerve to cortex. Single-unit recordings from the inferior colliculus (IC) and auditory cortex of cats demonstrate that neurons respond to both lead and lag sounds at ISDs above behavioral echo thresholds, but the response to the lag is reduced at shorter ISDs, consistent with localization dominance. Here the influence of the relative locations of the leading and lagging sources on the PE was measured behaviorally in a psychophysical task and physiologically in the IC of awake behaving cats. At all configurations of lead-lag stimulus locations, the cats behaviorally exhibited summing localization, localization dominance, and breakdown of fusion. Recordings from the IC of awake behaving cats show neural responses paralleling behavioral measurements. Both behavioral and physiological results suggest systematically shorter echo thresholds when stimuli are further apart in space.

scholarly journals Behavior and modeling of two-dimensional precedence effect in head-unrestrained cats

2015 ◽  
Vol 114 (2) ◽  
pp. 1272-1285
Author(s):  
Yan Gai ◽  
Janet L. Ruhland ◽  
Tom C. T. Yin
Keyword(s):  
Potassium Current ◽  
Horizontal Plane ◽  
Sagittal Plane ◽  
Precedence Effect ◽  
The Novel ◽  
Medial Superior Olive ◽  
Localization Dominance ◽  
Low Threshold ◽  
Spatial Locations ◽  
Auditory Illusion

The precedence effect (PE) is an auditory illusion that occurs when listeners localize nearly coincident and similar sounds from different spatial locations, such as a direct sound and its echo. It has mostly been studied in humans and animals with immobile heads in the horizontal plane; speaker pairs were often symmetrically located in the frontal hemifield. The present study examined the PE in head-unrestrained cats for a variety of paired-sound conditions along the horizontal, vertical, and diagonal axes. Cats were trained with operant conditioning to direct their gaze to the perceived sound location. Stereotypical PE-like behaviors were observed for speaker pairs placed in azimuth or diagonally in the frontal hemifield as the interstimulus delay was varied. For speaker pairs in the median sagittal plane, no clear PE-like behavior occurred. Interestingly, when speakers were placed diagonally in front of the cat, certain PE-like behavior emerged along the vertical dimension. However, PE-like behavior was not observed when both speakers were located in the left hemifield. A Hodgkin-Huxley model was used to simulate responses of neurons in the medial superior olive (MSO) to sound pairs in azimuth. The novel simulation incorporated a low-threshold potassium current and frequency mismatches to generate internal delays. The model exhibited distinct PE-like behavior, such as summing localization and localization dominance. The simulation indicated that certain encoding of the PE could have occurred before information reaches the inferior colliculus, and MSO neurons with binaural inputs having mismatched characteristic frequencies may play an important role.

A Neuronal Correlate of the Precedence Effect Is Associated With Spatial Selectivity in the Barn Owl's Auditory Midbrain

2004 ◽  
Vol 92 (4) ◽  
pp. 2051-2070 ◽  
Author(s):  
Matthew W. Spitzer ◽  
Avinash D. S. Bala ◽  
Terry T. Takahashi
Keyword(s):  
Receptive Field ◽  
Inferior Colliculus ◽  
Sound Localization ◽  
Auditory Processing ◽  
Precedence Effect ◽  
Auditory Midbrain ◽  
Spatial Selectivity ◽  
Neuronal Correlate ◽  
Spatial Tuning ◽  
Underlying Mechanisms

Sound localization in echoic conditions depends on a precedence effect (PE), in which the first arriving sound dominates the perceived location of later reflections. Previous studies have demonstrated neurophysiological correlates of the PE in several species, but the underlying mechanisms remain unknown. The present study documents responses of space-specific neurons in the barn owl's inferior colliculus (IC) to stimuli simulating direct sounds and reflections that overlap in time at the listener's ears. Responses to 100-ms noises with lead-lag delays from 1 to 100 ms were recorded from neurons in the space-mapped subdivisions of IC in anesthetized owls (N2O/isofluorane). Responses to a target located at a unit's best location were usually suppressed by a masker located outside the excitatory portion of the spatial receptive field. The least spatially selective units exhibited temporally symmetric effects, in that the amount of suppression was the same whether the masker led or lagged. Such effects mirror the alteration of localization cues caused by acoustic superposition of leading and lagging sounds. In more spatially selective units, the suppression was often temporally asymmetric, being more pronounced when the masker led. The masker often evoked small changes in spatial tuning that were not related to the magnitude of suppressive effects. The association of temporally asymmetric suppression with spatial selectivity suggests that this property emerges within IC, and not at earlier stages of auditory processing. Asymmetric suppression reduces the ability of highly spatially selective neurons to encode the location of lagging sounds, providing a possible basis for the PE.

scholarly journals Information conveyed by inferior colliculus neurons about stimuli with aligned and misaligned sound localization cues

2011 ◽  
Vol 106 (2) ◽  
pp. 974-985 ◽  
Author(s):  
Sean J. Slee ◽  
Eric D. Young
Keyword(s):  
Inferior Colliculus ◽  
Sound Localization ◽  
Virtual Space ◽  
Central Nucleus ◽  
Single Neurons ◽  
Interaural Time Differences ◽  
Marmoset Monkey ◽  
Interaural Level Differences ◽  
Broadband Sound ◽  
Response Area

Previous studies have demonstrated that single neurons in the central nucleus of the inferior colliculus (ICC) are sensitive to multiple sound localization cues. We investigated the hypothesis that ICC neurons are specialized to encode multiple sound localization cues that are aligned in space (as would naturally occur from a single broadband sound source). Sound localization cues including interaural time differences (ITDs), interaural level differences (ILDs), and spectral shapes (SSs) were measured in a marmoset monkey. Virtual space methods were used to generate stimuli with aligned and misaligned combinations of cues while recording in the ICC of the same monkey. Mutual information (MI) between spike rates and stimuli for aligned versus misaligned cues were compared. Neurons with best frequencies (BFs) less than ∼11 kHz mostly encoded information about a single sound localization cue, ITD or ILD depending on frequency, consistent with the dominance of ear acoustics by either ITD or ILD at those frequencies. Most neurons with BFs >11 kHz encoded information about multiple sound localization cues, usually ILD and SS, and were sensitive to their alignment. In some neurons MI between stimuli and spike responses was greater for aligned cues, while in others it was greater for misaligned cues. If SS cues were shifted to lower frequencies in the virtual space stimuli, a similar result was found for neurons with BFs <11 kHz, showing that the cue interaction reflects the spectra of the stimuli and not a specialization for representing SS cues. In general the results show that ICC neurons are sensitive to multiple localization cues if they are simultaneously present in the frequency response area of the neuron. However, the representation is diffuse in that there is not a specialization in the ICC for encoding aligned sound localization cues.

scholarly journals Short-Latency, Goal-Directed Movements of the Pinnae to Sounds That Produce Auditory Spatial Illusions

2010 ◽  
Vol 103 (1) ◽  
pp. 446-457 ◽  
Author(s):  
Daniel J. Tollin ◽  
Elizabeth M. McClaine ◽  
Tom C. T. Yin
Keyword(s):  
Operant Conditioning ◽  
Sound Source ◽  
Neural Mechanisms ◽  
Precedence Effect ◽  
Eye Position ◽  
Sound Sources ◽  
The Gaze ◽  
Localization Dominance ◽  
Echo Threshold ◽  
Behavioral Constraint

The precedence effect (PE) is an auditory spatial illusion whereby two identical sounds presented from two separate locations with a delay between them are perceived as a fused single sound source whose position depends on the value of the delay. By training cats using operant conditioning to look at sound sources, we have previously shown that cats experience the PE similarly to humans. For delays less than ±400 μs, cats exhibit summing localization, the perception of a “phantom” sound located between the sources. Consistent with localization dominance, for delays from 400 μs to ∼10 ms, cats orient toward the leading source location only, with little influence of the lagging source. Finally, echo threshold was reached for delays >10 ms, where cats first began to orient to the lagging source. It has been hypothesized by some that the neural mechanisms that produce facets of the PE, such as localization dominance and echo threshold, must likely occur at cortical levels. To test this hypothesis, we measured both pinnae position, which were not under any behavioral constraint, and eye position in cats and found that the pinnae orientations to stimuli that produce each of the three phases of the PE illusion was similar to the gaze responses. Although both eye and pinnae movements behaved in a manner that reflected the PE, because the pinnae moved with strikingly short latencies (∼30 ms), these data suggest a subcortical basis for the PE and that the cortex is not likely to be directly involved.

Psychophysical and Physiological Evidence for a Precedence Effect in the Median Sagittal Plane

1997 ◽  
Vol 77 (4) ◽  
pp. 2223-2226 ◽  
Author(s):  
Ruth Y. Litovsky ◽  
Brad Rakerd ◽  
Tom C. T. Yin ◽  
William M. Hartmann
Keyword(s):  
Inferior Colliculus ◽  
Sagittal Plane ◽  
Precedence Effect ◽  
Single Neurons ◽  
Spectral Cues ◽  
Physiological Evidence ◽  
True Location

Litovsky, Ruth Y., Brad Rakerd, Tom C. T. Yin, and William M. Hartmann. Psychophysical and physiological evidence for a precedence effect in the median sagittal plane. J. Neurophysiol. 77: 2223–2226, 1997. A listener in a room is exposed to multiple versions of any acoustical event, coming from many different directions in space. The precedence effect is thought to discount the reflected sounds in the computation of location, so that a listener perceives the source near its true location. According to most auditory theories, the precedence effect is mediated by binaural differences. This report presents evidence that the precedence effect operates in the median sagittal plane, where binaural differences are virtually absent and where spectral cues provide information regarding the location of sounds. Parallel studies were conducted in psychophysics by measuring human listeners' performance, and in neurophysiology by measuring responses of single neurons in the inferior colliculus of cats. In both experiments the precedence effect was found to operate similarly in the azimuthal and sagittal planes. It is concluded that precedence is mediated by binaurally based and spectrally based localization cues in the azimuthal and sagittal planes, respectively. Thus,models that attribute the precedence effect entirely to processes that involve binaural differences are no longer viable.

Psychophysical Investigation of an Auditory Spatial Illusion in Cats: The Precedence Effect

2003 ◽  
Vol 90 (4) ◽  
pp. 2149-2162 ◽  
Author(s):  
Daniel J. Tollin ◽  
Tom C.T. Yin
Keyword(s):  
Horizontal Plane ◽  
Sagittal Plane ◽  
Source Location ◽  
Precedence Effect ◽  
Search Coil ◽  
Localization Dominance ◽  
Reverberant Environments ◽  
Echo Threshold ◽  
Search Coil Technique

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 μs 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 μs 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.

Neural responses to simple simulated echoes in the auditory brain stem of the unanesthetized rabbit

1995 ◽  
Vol 74 (6) ◽  
pp. 2469-2486 ◽  
Author(s):  
D. C. Fitzpatrick ◽  
S. Kuwada ◽  
R. Batra ◽  
C. Trahiotis
Keyword(s):  
Time Course ◽  
Superior Olivary Complex ◽  
Precedence Effect ◽  
Natural Environments ◽  
Complete Suppression ◽  
Sound Waves ◽  
Neural Basis ◽  
Unanesthetized Rabbit ◽  
Echo Threshold ◽  
Time Required

1. In most natural environments, sound waves from a single source will reach a listener through both direct and reflected paths. Sound traveling the direct path arrives first, and determines the perceived location of the source despite the presence of reflections from many different locations. This phenomenon is called the "law of the first wavefront" or "precedence effect." The time at which the reflection is first perceived as a separately localizable sound defines the end of the precedence window and is called "echo threshold." The precedence effect represents an important property of the auditory system, the neural basis for which has only recently begun to be examined. Here we report the responses of single neurons in the inferior colliculus (IC) and superior olivary complex (SOC) of the unanesthetized rabbit to a sound and its simulated reflection. 2. Stimuli were pairs of monaural or binaural clicks delivered through earphones. The leading click, or conditioner, simulated a direct sound, and the lagging click, or probe, simulated a reflection. Interaural time differences (ITDs) were introduced in the binaural conditioners and probes to adjust their simulated locations. The probe was always set at the neuron's best ITD, whereas the conditioner was set at the neuron's best ITD or its worst ITD. To measure the time course of the effects of the conditioner on the probe, we examined the response to the probe as a function of the conditioner-probe interval (CPI). 3. When IC neurons were tested with conditioners and probes set at the neuron's best ITD, the response to the probe as a function of CPI had one of two forms: early-low or early-high. In early-low neurons the response to the probe was initially suppressed but recovered monotonically at longer CPIs. Early-high neurons showed a nonmonotonic recovery pattern. In these neurons the maximal suppression did not occur at the shortest CPIs, but rather after a period of less suppression. Beyond this point, recovery was similar to that of early-low neurons. The presence of early-high neurons meant that the overall population was never entirely suppressed, even at short CPIs. Taken as a whole. CPIs for 50% recovery of the response to the probe among neurons ranged from 1 to 64 ms with a median of approximately 6 ms. 4. The above results are consistent with the time course of the precedence effect for the following reasons. 1) The lack of complete suppression at any CPI is compatible with behavioral results that show the presence of a probe can be detected even at short CPIs when it is not separately localizable. 2) At a CPI corresponding to echo threshold for human listeners (approximately 4 ms CPI) there was a considerable response to the probe, consistent with it being heard as a separately localizable sound at this CPI. 3) Full recovery for all neurons required a period much longer than that associated with the precedence effect. This is consistent with the relatively long time required for conditioners and probes to be heard with equal loudness. 5. Conditioners with either the best ITD or worst ITD were used to determine the effect of ITD on the response to the probe. The relative amounts of suppression caused by the two ITDs varied among neurons. Some neurons were suppressed about equally by both types of conditioners, others were suppressed more by a conditioner with the best ITD, and still others by a conditioner with the worst ITD. Because the best ITD and worst ITD presumably activate different pathways, these results suggest that different neurons receive a different balance of inhibition from different sources. 6. The recovery functions of neurons not sensitive to ITDs were similar to those of ITD-sensitive, neurons. This suggests that the time course of suppression may be common among different IC populations. 7. We also studied neurons in the SOC. Although many showed binaural interactions, none were sensitive to ITDs. Thus the response of this population may not be

Sign in / Sign up

Export Citation Format

Share Document