Localization of tones by horses: Use of binaural cues and the role of the superior olivary complex.

Assessed the ability of horses to use binaural time and intensity difference cues to localize sound in free-field localization test using pure tones. Ss were required to discriminate the locus of a single tone pip ranging in frequency from 250 Hz to 25 kHz emitted by loudspeakers located 30° to the left and right of the Ss' midline. Three Ss were tested with a 2-choice procedure; 2 additional Ss were tested with a conditioned avoidance procedure. Ss were able to localize 250 Hz, 500 Hz, and 1 kHz but not 2 kHz and above. Because the frequency of ambiguity for the binaural phase-difference cue was calculated to be 1.5 kHz, results indicate that Ss could use binaural time differences but not binaural intensity differences. This finding was supported by an unconditioned orientation test involving 4 additional Ss, who correctly oriented to a 500-Hz tone pip but not to an 8-kHz tone pip. Analysis of the superior olivary complex, the brain-stem nucleus at which binaural interactions first take place, reveals that the lateral superior olive (LSO) is relatively small in the horse and lacks the laminar arrangement of bipolar cells characteristic of the LSO of most mammals that can use binaural intensity differences. (36 ref) (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

Free-field binaural unmasking in ferrets.

Free-field detection by normal and monaural ferrets (N?=?4) of a 500-Hz tone presented over 1 laterally placed loudspeaker and partially masked by narrowband noise from 2 sources was studied at 2 angular separations of the noise sources (0° and 180°). Monaural listening was achieved either by plugging 1 ear canal or removing 1 cochlea. Normal ferrets showed an improvement in detectability of the tone when there was a 180° separation between the noise sources. This unmasking of the tone was abolished in both groups of monaural ferrets, suggesting that the unmasking was due to binaural processing. The development of an animal model demonstrating free-field binaural unmasking, in a species other than humans, will allow investigation into the functional consequences of experimental hearing loss. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Dichotic pitches as illusions of binaural unmasking. I. Huggins' pitch and the "binaural edge pitch"

The two most salient dichotic pitches, the Huggins pitch (HP) and the binaural edge pitch (BEP), are produced by applying interaural phase transitions of 360 and 180 degrees, respectively, to a broadband noise. This paper examines accounts of these pitches, concentrating on a "central activity pattern" (CAP) model and a "modified equalization-cancellation" (mE-C) model. The CAP model proposes that a dichotic pitch is heard at frequency f when an individual across-frequency scan in an interaural cross-correlation matrix contains a sharp peak at f. The mE-C model proposes that a dichotic pitch is heard when a plot of interaural decorrelation against frequency contains a peak at f. The predictions of the models diverge for the BEP at very narrow transition bandwidths: the mE-C model predicts that salience is sustained, while the CAP model predicts that salience declines and that the dominant percept is of the in-phase segment of the noise. Experiment 1 showed that the salience of the BEP was sustained at the narrowest bandwidths that could be generated (0.5% of the transition frequency). Experiment 2 confirmed that the pitch of a BEP produced by a 0.5% transition bandwidth was close to the frequency of the transition band. Experiment 3 showed that pairs of simultaneous narrow 180-degree transitions, whose frequencies corresponded to vowel formants, were perceived as the intended vowels. Moreover, the same vowels were perceived whether the in-phase portion of the noise lay between the two transition frequencies or on either side of them. In contrast, different patterns of identification responses were made to diotic band-pass and band-stop noises whose cutoff frequencies corresponded to the same formants. Thus, the vowel-identification responses made to the dichotic stimuli were not based on hearing the in-phase portions of the noise as formants. These results are not predicted by the CAP model but are consistent with the mE-C model. It is argued that the mE-C model provides a more coherent and parsimonious account of many aspects of the HP and the BEP than do alternative models.     

Interaural delay-dependent changes in the binaural difference potential in cat auditory brainstem response: implications about the origin of the binaural interaction component

Auditory brainstem responses (ABRs) evoked by dichotic clicks with 12 different interaural delays (ITDs) between 0 and 1500 microsecond(s) were recorded from the vertices of 10 cats under ketamine anesthesia. The so-called binaural difference potential (BDP), considered to be an indicator of binaural interaction (BI), was computed by subtracting the sum of the two monaural responses from the binaural one. The earliest and most prominent component of BDP was a negative deflection (DN1) at a latency between 4 and 4.8 ms. Like all the other components of BDP, DNI was also due to binaural reduction rather than enhancement of the corresponding ABR wave, P4 in this case. Furthermore, the way its latency increased as a function of ITD was also not compatible with what would be predicted by the delay-line coincidence detector models based on the excitatory-excitatory units in the medial superior olive (MSO). We therefore proposed an alternative hypothesis for the origin of this BI component based on the inhibitory-excitatory (IE) units in the lateral superior olive (LSO). The computational model designed closely simulated the ITD-dependent attenuation and latency shifts observed in DN1. It was therefore concluded that the origin of this BI component in the cat's vertex-ABR could be the lateral lemniscal output of the LSO, although the delay lines which have been shown to exist also in the mammalian brain may play an important role in encoding ITDs.     

Receptive musical aptitude using a binaural frequency discrimination test

Musical aptitude is the most important precondition for any applicant taking up studies at a music conservatory. Thus, the applicants are tested and assessed very thoroughly by experienced music teachers. To support these subjective results by objective data a procedure was developed allowing to test both the examinee's frequency and octave discrimination capacity in a simple way. In a 1-Hz cycle the frequencies f1 and f2 are directed mutually to the right and left ear, respectively, at a defined sound level. Then the examinee is asked to syntonize e.g. frequency equality f1 = f2 (or f1 = 2 x f2). It can be shown that there is a direct correlation between the amount of deviation from nominal frequency and subjective assessment of musicality.     

Monaural and binaural story recall by schizophrenic subjects.

P. Green and other investigators have reported that schizophrenic Ss have poorer recall of stories presented to both ears than to the single best ear (binaural deficit) and poorer recall of stories presented to the left ear than to the right ear (monaural asymmetry) than do normal control Ss. These studies are plagued by potential methodological problems, including differences in overall accuracy, which artifactually affect the difference scores, and scoring methods that are vulnerable to systematic bias. In this study, scores of schizophrenic, bipolar, and normal control Ss on the Auditory Comprehension Test were compared. Scoring bias was avoided by the use of blind scoring and a revised scoring manual, and artifactual effects of accuracy were considered in interpreting the results. Contrary to previous findings, the groups did not differ on either monaural asymmetry or binaural deficit. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The effects of attenuation of frequency segments on binaural localization of sound

Perceived location of tonal stimuli d and narrow noise bands presented in two-dimensional space varies in an orderly manner with changes in stimulus frequency. Hence, frequency has a referent in space that is most apparent during monaural listening. The assumption underlying the present study is that maximum sound pressure level measured at the ear canal entrance for the various frequencies serves as a prominent spectral cue for their spatial referents. Even in binaural localization, location judgments in the vertical plane are strongly influenced by spatial referents. We measured sound pressure levels at the left ear canal entrance for 1.0-kHz-wide noise bands, centered from 4.0 kHz through 10.0 kHz, presented at locations from 60 degrees through -45 degrees in the vertical plane; the horizontal plane coordinate was fixed at -90 degrees. On the basis of these measurements, we fabricated three different bandstop stimuli in which differently centered 2.0-kHz-wide frequency segments were filtered from a broadband noise. Unfiltered broadband noise served as the remaining stimulus. Localization accuracy differed significantly among stimulus conditions (p < .01). Where in the vertical plane most errors were made depended on which frequency segment was filtered from the broadband noise.     

Free-field binaural unmasking in budgerigars ( Melopsittacus undulatus ).

The detection of signals in noise is important for understanding both the mechanisms of hearing and how the auditory system functions under more natural conditions. In humans, the auditory system gains some improvement if the signal and noise are separated in space (binaural masking release). Birds with small heads are at a disadvantage in separating noise and signal sources relative to large mammals, because interaural time differences are much smaller. Two binaural phenomena in budgerigars related to the detection of tones in noise were examined. Budgerigars show 8 dB of free-field binaural masking release when signal and noise are presented to their right side and correlated noise is presented to their left side. Budgerigars also show a spatial masking release of 9 dB when a signal and noise are separated in azimuth by 90°. These results are similar to those found in humans and other mammals with much larger heads. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The normalized correlation: accounting for binaural detection across center frequency

Bernstein and Trahiotis [L. R. Bernstein and C. Trahiotis, J. Acoust. Soc. Am. 100, 1754-1763 (1996)] recently reported the results of experiments designed to determine the form of interaural correlation that accounts for listeners' sensitivities to interaural disparities within high-frequency stimuli. Overall, those results demonstrated that listeners' abilities to discriminate changes in the interaural correlation of the envelope (from a base correlation of 1.0) were well accounted for by the use of the normalized correlation. The purpose of this study was to determine how well the normalized correlation computed subsequent to half-wave rectification and low-pass filtering could account for binaural detection data at low, intermediate, and high frequencies, respectively. In a four-interval, two-alternative task, listeners detected which interval contained a tone (between 500 Hz and 2 kHz) added antiphasically to diotic, 100-Hz-wide, noise (NoS pi). "Nonsignal" intervals contained the tone added homophasically (NoSo). Performance was measured for signal-to-noise ratios between -30 and +30 dB. Results indicated that a low-pass filter function based on physiological measures of synchrony in cochlear nerve fibers in conjunction with the assumption of half-wave, square-law rectification, accounted for typically 80% of the variance in the behavioral data.     

An auditory illusion predicted from a weighted cross-correlation model of binaural interaction.

In humans, the lateral movement of an acoustic source produces dynamic changes in the relative sound-pressure level and time of arrival of the acoustic wave at the 2 ears. The dynamic nature of these cues is assumed to play an important role in the perception of lateral motion. A phenomenon of auditory motion is reported whose lateral direction and relative velocity may be specified while interaural differences are kept constant. The stimulus producing this percept is a narrowband waveform whose instantaneous bandwidth is a cosine function of time. This phenomenon is predicted from a model of cross-correlation that estimates the running position of an image from a weighted combination of 2 variables: (a) magnitude of interaural delay, with smaller delays receiving more weight, and (b) consistency of interaural information across frequency. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Superior speech comprehension in schizophrenics under monaural versus binaural listening conditions.

20 schizophrenics, 10 nurses, and 10 psychiatric controls (all under age 60) listened to stories and answered the corresponding questions in 3 conditions. All task information was provided either binaurally or exclusively to the left or right ear, and scores were derived from the number of questions answered correctly in each condition. It was hypothesized that schizophrenics would display significant deficits in left ear speech comprehension on the assumption that the patients suffered from poor interhemispheric transfer, which had been observed on manual and visual tasks. Significant left ear deficits were observed in the schizophrenics but not in the controls. An unexpected effect, which may also reflect defective interhemispheric transfer, was that the schizophrenics, but not the controls, displayed significant deficits in binaural relative to right ear speech comprehension. It may be possible to increase the speech comprehension of schizophrenics who show this effect by a simple method that takes advantage of the observed right ear superiority. (30 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Temporal and binaural properties in dorsal cochlear nucleus and its output tract

The dorsal cochlear nucleus (DCN) is one of three nuclei at the terminal zone of the auditory nerve. Axons of its projection neurons course via the dorsal acoustic stria (DAS) to the inferior colliculus (IC), where their signals are integrated with inputs from various other sources. The DCN presumably conveys sensitivity to spectral features, and it has been hypothesized that it plays a role in sound localization based on pinna cues. To account for its remarkable spectral properties, a DCN circuit scheme was developed in which three inputs converge onto projection neurons: auditory nerve fibers, inhibitory interneurons, and wide-band inhibitors, which possibly consist of Onset-chopper (Oc) cells. We studied temporal and binaural properties in DCN and DAS and examined whether the temporal properties are consistent with the model circuit. Interneurons (type II) and projection (types III and IV) neurons differed from Oc cells by their longer latencies and temporally nonlinear responses to amplitude-modulated tones. They also showed evidence of early inhibition to clicks. All projection neurons examined were inhibited by stimulation of the contralateral ear, particularly by broadband noise, and this inhibition also had short latency. Because Oc cells had short-latency responses and were well driven by broadband stimuli, we propose that they provide short-latency inhibition to DCN for both ipsilateral and contralateral stimuli. These results indicate more complex temporal behavior in DCN than has previously been emphasized, but they are consistent with the recently described nonlinear behavior to spectral manipulations and with the connectivity scheme deduced from such manipulations.     

Detectability index measures of binaural masking level difference across populations of inferior colliculus neurons

In everyday life we continually need to detect signals against a background of interfering noise (the "cocktail party effect"): a task that is much easier to accomplish using two ears. The binaural masking level difference (BMLD) measures the ability of listeners to use a difference in binaural attributes to segregate sound sources and thus improve their discriminability against interfering noises. By computing the detectability of tones from rate-versus-level functions in the presence of a suprathreshold noise, we previously demonstrated that individual low-frequency delay-sensitive neurons in the inferior colliculus are able to show BMLDs. Here we consider the responses of a population of such neurons when the noise level is held constant (as conventionally in psychophysical paradigms). We have sampled the responses of 121 units in the inferior colliculi of five guinea pigs to identical noise and 500 Hz tones at both ears (NoSo) and to identical noise but with the 500 Hz tone at one ear inverted (NoSpi). The result suggests that the neurons subserving detection of So tones in No (identical noise at the two ears) noise are those neurons with best frequencies (BFs) close to 500 Hz that respond to So tones with an increase in their discharge rate from that attributable to the noise. The detection of the inverted (Spi) signal is also attributable to neurons with BFs close to 500 Hz. However, among these neurons, the presence of the Spi tone was indicated by an increased discharge rate in some neurons and by a decreased discharge rate in others.     

Neuroanatomical basis of binaural phase-difference analysis for sound localization: A comparative study.

4 varieties of mammals whose medial superior olives range from large to none at all were tested for their ability to localize single, brief tone pips at various frequencies. Ss were 2 hedgehogs, 2 Charles River white rats, 2 tree shrews, and 1 cat. Although each animal could localize high-frequency tone pips, their ability to localize middle- and low-frequency tone pips corresponded to the size of their medial superior olive (MSO). Since this latter range of frequencies is the one in which binaural phase-difference cues predominate, this anatomical-behavioral correspondence supports the idea that MSO is the chief binaural time analyzing center for sound localization. (19 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Localization of noise, use of binaural cues, and a description of the superior olivary complex in the smallest carnivore, the least weasel (Mustela nivalis).

Cats and dogs have relatively good sound-localization acuity, and the question arises as to whether this trait is a characteristic of all carnivores or whether it is due to the fact that they have large heads and correspondingly large binaural localization cues available to them. The localization acuity of the least weasel, the smallest extant carnivore, was found to be less accurate than larger carnivores but more accurate than other small mammals. This suggests that carnivores may be under strong selective pressure to localize accurately but that interaural distance may be a limiting factor. The least weasel is capable of using both binaural phase differences and intensity differences to localize, but has a relatively broad mid-frequency range for which neither cue is optimal. Finally, the superior olivary complex of the least weasel is well developed and resembles that of larger carnivores more than that of small rodents. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Model system of binaural hearing. III. Binaural processing of information about short sounds

A model of neuronal structures of the medial superior olivary nuclei and inferior colliculus performing the determination of the direction on the source of the short (shorter than 10 ms) sound signals is presented. In the model the difference between the moments of the arrival of the two informational messages formed on the stage of monaural information processing is calculated. The result of this calculation is the firing probability of the primary detector (the neuron of the medial superior olivary nucleus). Because of the internal noise the curve of this probability as a function of the direction on the source is smoothly sloped. The estimation of the direction is the result of the statistical processing of the responses of the primary detectors ensemble. The direction on the source of sound is coded by a position of the secondary detector (the neuron of the inferior colliculus) on the direction "scale".