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This paper discusses how the barn owl's brain stem auditory pathway is divided into two physiologically and anatomically segregated channels for separate processing of interaural phase and intensity cues for sound localization. The paper also points out the power of the "downstream" approach by which the emergence of a higher-order neuron's stimulus selectivity can be traced through lower-order stations.
Direct electrical stimulation of the auditory nerve can be used to restore some degree of hearing to the profoundly deaf. Percepts due to electrical stimulation have characteristics corresponding approximately to the acoustic percepts of loudness, pitch, and timbre. To encode speech as a pattern of electrical stimulation, it is necessary to determine the effects of the stimulus parameters on these percepts. The effects of the three basic stimulus parameters of level, repetition rate, and stimulation location on subjects' percepts were examined. Pitch difference limens arising from changes in rate of stimulation increase as the stimulating rate increases, up to a saturation point of between 200 and 1000 pulses per second. Changes in pitch due to electrode selection depend upon the subject, but generally agree with a tonotopic organization of the human cochlea. Further, the discriminability of such place-pitch percepts seems to be dependent on the degree of current spread in the cochlea. The effect of stimulus level on perceived pitch is significant but is highly dependent on the individual tested. The results of these experiments are discussed in terms of their impact on speech-processing strategies and their relevance to acoustic pitch perception.
Auditory event-related brain potentials (ERPs) and reaction times were analyzed in a selective attention task in which subjects attended to tone pips presented at high rates (interstimulus intervals [ISIs] of 40-200 msec). Subjects responded to infrequent target tones of a specified frequency (250 or 4000 Hz) and location (left or right ear) that were louder than otherwise identical tones presented randomly to the left and right ears. Negative difference (Nd) waves were isolated by subtracting ERPs to tones with no target features from ERPs to the same tones when they shared target location, frequency, or both frequency and location cues. Nd waves began 60-70 msec after tone onset and lasted until 250-350 msec after tone onset, even for tones with single attended cues. The duration of Nd waves exceeded the ISIs between successive tones, implying that several stimuli underwent concurrent analysis. Nd waves associated with frequency processing had scalp distributions different from those associated with location processing, implying that the features were analyzed in distinct cortical areas. Nd waves specific to auditory feature conjunction were isolated. These began at latencies of 110-120 msec, some 30-40 msec after the Nds to single features. The relative timing of the different Nd waves suggests that auditory feature conjunction begins after a brief parallel analysis of individual features but before feature analysis is complete.
Previous study demonstrated that responsiveness to electrical stimulation of subcortical areas of cats was enhanced if preceded by a tone pip. Habituation occurred when the tone was repeatedly presented. In this study, the capacity for generalization of a habituated tone was measured by occasionally interpolating a tone of different frequency or intensity among a group of monotonously presented standard tones, measuring responsiveness in each case. The responses were: (1) the visual cortical evoked potential, (2) evoked EMG activity, and (3) evoked head movement. Frequency shifts and intensity increases produced recovery of responsiveness at latencies as short as 90 msec, while decreases had no effect on recovery. Responsiveness increased with degree of frequency shift up to 576 Hz, but showed no further increase for greater shifts up to 2304 Hz. A shift of 16 Hz was effective, comparable to behavioral thresholds for frequency discrimination. Thus, these gradients agreed with the form predicted on the basis of central fatigue models of habituation and the bandwidth of the eighth nerve fibers.
A brief, vivid phase of auditory sensory storage that outlasts the stimulus could be used in perception in two ways: First, all of the neural activity resulting from the stimulus, including that of the sensory store, could contribute to a sensation of growing loudness; second, the sensory store could permit the continued extraction of information about the sound's acoustic properties. This study includes a task for which these two processes lead to different predictions; a third prediction is based on the two processes combined. The task required loudness judgments for two brief tones presented with a variable intertone interval. The results of Experiments 1-3 were as one would expect if both the growth of sensation and information extraction contributed to the pattern of loudness judgments. Experiment 4 strengthened the two-process account by demonstrating the separability of the two processes. Approaches to mathematical modeling of these results are discussed.
This study investigated the effect of intensity on pitch in electric hearing and its relationship to the speech perception ability of cochlear implantees. Subjects were 13 adult users of the Nucleus 22 cochlear implant system, using either the Spectra22 or ESPrit22 speech processor and the SPEAK speech processing strategy. A multidimensional scaling technique was employed. Speech perception was measured using sentences and vowels. All measurements were performed in a soundfield condition, and subjects wore their own speech processors with their normally used settings. Results showed a significant correlation between the degree of deviation of the subjects' stimulus spaces from the "ideal" space and subjects' performance with the sentences, but not with the vowels. A significant correlation was found between subjects' response variability in performing the multidimensional scaling task and their speech perception measures, suggesting that spectral smearing or underlying cognitive abilities might affect implantees' speech perception performance.
To study the regional cerebral blood flow (rCBF) correlates of auditory processing, we performed rCBF measurements in young, normal right-handed volunteers engaged in listening tasks. Using the xenon Xe 133 inhalation technique, rCBF was measured in 11 regions in each hemisphere. Compared with a baseline condition, significant rCBF increases in the left posterior Sylvian regions were associated with both verbal phonologic processing and nonverbal acoustic processing. Verbal semantic processing was associated with unilateral increased rCBF in the posterior Sylvian region contralateral to the hand used for response signaling. Comparison of rCBF from homologous regions of the two hemispheres also confirmed a clear difference between the rhyme detection and meaning detection tasks, with the former strongly lateralized to the left hemisphere. Thus, changes in rCBF were related to the nature of the listening task.
Single neurones in the central nucleus of the inferior colliculus (ICC) of barbiturate-anesthetized cats were examined using free-field, pure-tone stimuli of low intensity at the neurones' best frequency. Receptive field size was inversely correlated with best frequency. Almost all neurones were maximally excited by stimulus positions in the hemifield contralateral to the recording electrode, irrespective of their best frequency. Simultaneous cochlear microphonic recording revealed that the neurones' best excitatory area was also the spatial region associated with maximum amplification by the contralateral outer ear. This amplification resulted in extremely low (less than -20 dB SPL in some neurones) best frequency thresholds. Response patterns were found not to vary markedly with speaker position. The results suggest that most ICC neurones are more sensitive to stimulation of the contralateral ear than to stimulation of the ipsilateral ear.
This study aimed to classify the responses of single units in the auditory midbrain to acoustic stimuli presented in the free field in order to characterize those units likely to have a role in sound localization in the horizontal plane. The responses of 131 single units in the inferior colliculus of the cat and the brush-tailed possum were studied using tone and noise-burst stimuli presented from a speaker capable of movement at any point along a plane 10 degrees above the horizontal plane. Speaker positions along this plane are referred to as speaker azimuths; those on the same side as the recorded inferior colliculus as ipsilateral, and on the opposite side as contralateral, azimuths. For each unit, spike counts were measured as a function of azimuth either at the best frequency (BF) or using noise bursts. These functions are referred to as azimuth functions and were usually measured for at least two intensities, between 10 and 70 dB above threshold. The recording sites of most units were identified histologically with the aid of microlesions and were related to the major subdivisions of the inferior colliculus: the central nucleus (ICC), the lateral part of the external nucleus (ICX), and the rostroventral process (R-ICX). Two units were located in the pericentral nucleus and two in the dorsal nucleus of the lateral lemniscus. Two major classes of neuron were identified: omnidirectional and directionally sensitive. Omnidirectional units exhibited azimuth functions that were either flat or that declined gradually at progressively ipsilateral azimuths. For the latter units, discharge rates at all points monotonically increased with stimulus intensity. There was no indication, for either type of omnidirectional unit, of significant binaural interaction. A good correlation was found between the summed proportions of excitatory-excitatory (EE) and monaural (EO) units observed in dichotic studies (46-55%) and the proportion of omnidirectional units in the present study (47%). A subgroup of directionally sensitive units (36% of the total) displayed azimuth functions for which the azimuthal position of the discharge border or peak firing azimuth remained essentially unaltered over a range of stimulus intensities. These azimuth-selective units are likely to have a role in the detection of the location of stimuli in the horizontal plane and appear to include units that would be considered excitatory-inhibitory (EI) or delay sensitive in dichotic studies. The azimuths over which directionally sensitive units showed their marked directional effects were influenced by the position of the contralateral pinna.(ABSTRACT TRUNCATED AT 400 WORDS)
Historically, the primary concern for hearing aid design and fitting is optimization for speech inputs. However, increasingly other types of inputs are being investigated and this is certainly the case for music. Whether the hearing aid wearer is a musician or merely someone who likes to listen to music, the electronic and electro-acoustic parameters described can be optimized for music as well as for speech. That is, a hearing aid optimally set for music can be optimally set for speech, even though the converse is not necessarily true. Similarities and differences between speech and music as inputs to a hearing aid are described. Many of these lead to the specification of a set of optimal electro-acoustic characteristics. Parameters such as the peak input-limiting level, compression issues-both compression ratio and knee-points-and number of channels all can deleteriously affect music perception through hearing aids. In other cases, it is not clear how to set other parameters such as noise reduction and feedback control mechanisms. Regardless of the existence of a "music program,'' unless the various electro-acoustic parameters are available in a hearing aid, music fidelity will almost always be less than optimal. There are many unanswered questions and hypotheses in this area. Future research by engineers, researchers, clinicians, and musicians will aid in the clarification of these questions and their ultimate solutions.
The acceptance of cochlear implantation as an effective and safe treatment for deafness has increased steadily over the past quarter century. The earliest devices were the first implanted prostheses found to be successful in compensating partially for lost sensory function by direct electrical stimulation of nerves. Initially, the main intention was to provide limited auditory sensations to people with profound or total sensorineural hearing impairment in both ears. Although the first cochlear implants aimed to provide patients with little more than awareness of environmental sounds and some cues to assist visual speech-reading, the technology has advanced rapidly. Currently, most people with modern cochlear implant systems can understand speech using the device alone, at least in favorable listening conditions. In recent years, an increasing research effort has been directed towards implant users' perception of nonspeech sounds, especially music. This paper reviews that research, discusses the published experimental results in terms of both psychophysical observations and device function, and concludes with some practical suggestions about how perception of music might be enhanced for implant recipients in the future. The most significant findings of past research are: (1) On average, implant users perceive rhythm about as well as listeners with normal hearing; (2) Even with technically sophisticated multiple-channel sound processors, recognition of melodies, especially without rhythmic or verbal cues, is poor, with performance at little better than chance levels for many implant users; (3) Perception of timbre, which is usually evaluated by experimental procedures that require subjects to identify musical instrument sounds, is generally unsatisfactory; (4) Implant users tend to rate the quality of musical sounds as less pleasant than listeners with normal hearing; (5) Auditory training programs that have been devised specifically to provide implant users with structured musical listening experience may improve the subjective acceptability of music that is heard through a prosthesis; (6) Pitch perception might be improved by designing innovative sound processors that use both temporal and spatial patterns of electric stimulation more effectively and precisely to overcome the inherent limitations of signal coding in existing implant systems; (7) For the growing population of implant recipients who have usable acoustic hearing, at least for low-frequency sounds, perception of music is likely to be much better with combined acoustic and electric stimulation than is typical for deaf people who rely solely on the hearing provided by their prostheses.
The purpose of this investigation was to determine what subjective factors may influence the success of fitting in-the-canal hearing instruments (ITCs). Four different types of ITC were fitted to four matched groups of 20 experienced ITC users. In addition to the subjective experiences of the users, the results were checked using insertion gain (IG) measurements. The results indicate that too sharp sound quality from paper rustle, from running water and from use of kitchen utensils is a common problem that it is important to be aware of. The possibility of maximal venting is of special importance in order to avoid autophony. Feedback may be a problem, especially when maximal venting is necessary. However, for several models of ITC, feedback is, in reality, a minor problem. High cost of batteries may be a problem for persons with ITC using 10/230 batteries. IG measurements indicated that a gain of approximately 0.3 in relation to the hearing losses was preferred. It may be advantageous to choose a model of ITC that has extensive flexibility.
The effects of cholinergic drugs on three different perceptual bisection tasks were studied in rats. Physostigmine (0.056-0.56 mg/kg), a reversible anticholinesterase, produced dose-dependent decrements in discriminability (A'), but did not affect the bisection point (BP) in visual duration, auditory duration, and auditory intensity bisection tasks. This finding is consistent with results previously obtained in an auditory duration bisection task with an irreversible anticholinesterase, diisopropyl phosphofluoridate. Scopolamine (0.075-0.422 mg/kg), a muscarinic cholinergic-receptor antagonist, produced dose-dependent decrements in both A' and BP in visual and auditory duration bisection tasks. The behavioral antagonism between physostigmine (0.56 mg/kg) and scopolamine (0.075-0.237 mg/kg) was studied in the visual and auditory duration bisection tasks. The BP was not affected by physostigmine alone or in combination with scopolamine, except at the largest dose of scopolamine, which produced a reliable decrement in the BP. A', however, was equally decreased by physostigmine alone and all combinations of physostigmine and scopolamine. Pirenzepine (1, 3 and 10 mg/kg), a selective high-affinity M1 muscarinic antagonist, had no effect on A' or the BP in the duration bisection tasks, suggesting changes in perception produced by muscarinic antagonists do not involve the M1 receptor subtype. The similar drug effects in different sensory modalities (visual and auditory) and perceptual systems (subjective duration and loudness) suggest that cholinergic drugs may affect perceptual mechanisms responsible for sensory coding, such as the output of a neural generator.
In a combined psychophysical-electrophysiological study on 29 patients with multiple sclerosis (MS), a compromised ability to make interaural time discriminations was nearly always found to be associated with 'abnormal' brain stem potentials evoked by clicks to at least one ear. However, no obvious relationships were found between evoked brain stem potentials and several other auditory behavioral measures such as interaural intensity discrimination, pure-tone thresholds and speech discrimination.
The amount of melanin pigmentation in the inner ear is positively correlated with the general pigmentation of the body and specifically with the amount of pigment in the eye. The misrouting of retinofugal projections which accompanies ocular and oculocutaneous albinism has been thought to be a defect in decussation unique to the visual system. Evidence suggests that functional abnormalities may also exist in the auditory systems of albino humans and animals. To evaluate this possibility, evoked potential techniques were used to examine the functional anatomy of decussating brainstem auditory pathways in albino and pigmented cats. Auditory brainstem responses (ABRs) were recorded from albino, pigmented, and Siamese cats using monaural stimulation. ABRs were recorded ipsilateral and contralateral to the stimulated ear. The albinos were complete tyrosinase-negative (cc), not the dominant white (W) variety associated with deafness. In pigmented cats, the amplitudes of ABRs recorded with the reference electrode ipsilateral to the stimulated ear and the ABRs recorded using the reference contralateral to the stimulated ear did not differ by more than 40% for individual components appearing between 2 and 4 ms after stimulus onset. In albino cats the components at these latencies were obliterated or greatly attenuated in the ABR recorded using the reference contralateral to the stimulated ear. These data indicate that anomalies may exist in the brainstem at the level of the acoustic striae, superior olivary nuclei and/or trapezoid body in tyrosinase-negative albino cats.
Patients with schizophrenia exhibit reduced levels of both prepulse inhibition of the startle reflex (PPI) and condition-test suppression of the P50 event-related potential. This study investigated the extent to which PPI and P50 suppression, which exhibit similar parametric sensitivities, are intrinsically auditory phenomena or can be induced cross-modally, and reflect common or distinct neural mechanisms of inhibition. PPI, N100, and P50 were assessed in 20 healthy male volunteers, using auditory test probes and both visual and auditory lead stimuli, separated by 100- or 500-ms interstimulus intervals (ISIs). PPI was found in the auditory-lead condition across the complete group, and with visual-lead stimuli in approximately half of the subjects. Intra-modal auditory PPI was significantly higher with the 100-ms ISI than with the 500-ms ISI. P50 suppression was found only with the 500-ms ISI, with no difference between the auditory and visual conditions. Source analyses revealed that suppression was associated with frontal cortical activity. N100 suppression was found only in the auditory condition, with no difference between 100- and 500-ms ISIs. Although both phenomena are considered to provide operational measures of gating, PPI and P50 suppression are differentially sensitive to ISI and therefore reflect partly different neural mechanisms. They are not intrinsically auditory phenomena, and both appear to involve frontal cortical activity. In contrast, N100 suppression is most likely based on refractory mechanisms intrinsic to the auditory system.
Habituation of human scalp-recorded cerebral evoked potentials was studied in response to auditory and visual repetitive stimuli of different intensities. Changes in magnitudes of evoked potentials with stimulus repetition were examined according to the parametric characteristics of habituation, generalization, and dishabituation. In addition, tests of the predictions of two theories of habituation were made regarding the degree and direction of intensity generalization of habituation. Both auditory and visual evoked potentials exhibited decrements in response magnitudes across the repetitive stimuli consistent with the parametric criteria of habituation. Early evoked potential peak components showed a pattern of intensity generalization of habituation consistent with the predictions of the dual-process theory of habituation. Intensity generalization of late evoked potential peak components occurred in a manner more consistent with the predictions of the stimulus comparator theory of habituation. These results provide further evidence that evoked potentials can be used as electrophysiological indexes of plasticity in humans.
The mechanical nonlinearity of the cochlea that is associated with normal cochlear function, induces distortion products that can be recorded in the external auditory canal. The acoustic cubic distortion product 2F1-F2 (DP) was measured in the external canal in the presence and in the absence of a contralateral white noise. The experiments were carried out on 20 guinea pigs after a section of the middle ear muscles. They showed that the presence of a contralateral white noise induces a significant reversible reduction of the magnitude of the DPs. This suppressive effect produced by the contralateral white noise was completely canceled out by the midline sagittal section of the brainstem. This report supports the hypothesis that the suppressive effect of a contralateral sound stimulation is mediated by the medial efferent system.