Psychoacoustics publications

Little is known about the neural bases of the reduced auditory and cortical processing speeds that have been recorded in language-impaired, autistic, schizophrenic, and other disabled human populations. Although there is strong evidence for genetic contributions to etiologies, epigenetic factors such as perinatal anoxia (PA) have been argued to be contributors, or causal, in a significant proportion of cases. In this article, we explored the consequences of PA on this elementary aspect of auditory behavior and on auditory system function in rats that were briefly perinatally anoxic. PA rats had increased acoustic thresholds and reduced processing efficiencies recorded in an auditory behavioral task. These rats had modestly increased interpeak intervals in their auditory brainstem responses, and substantially longer latencies in poststimulus time histogram responses recorded in the primary auditory cortex. The latter were associated with degraded primary auditory cortex receptive fields and a disrupted tonotopy. These processing deficits are consistent with the parallel behavioral and physiological deficits recorded in children and adults with a history of language-learning impairment and autism.

Using a closed-set speech recognition paradigm thought to be heavily influenced by informational masking, auditory selective attention was measured in 38 children (ages 4–16years) and 8 adults (ages 20–30years). The task required attention to a monaural target speech message that was presented with a time-synchronized distracter message in the same ear. In some conditions a second distracter message or a speech-shaped noise was presented to the other ear. Compared to adults, children required higher target∕distracter ratios to reach comparable performance levels, reflecting more informational masking in these listeners. Informational masking in most conditions was confirmed by the fact that a large proportion of the errors made by the listeners were contained in the distracter message(s). There was a monotonic age effect, such that even the children in the oldest age group (13.6–16years) demonstrated poorer performance than adults. For both children and adults, presentation of an additional distracter in the contralateral ear significantly reduced performance, even when the distracter messages were produced by a talker of different sex than the target talker. The results are consistent with earlier reports from pure-tone masking studies that informational masking effects are much larger in children than in adults.

We hypothesized that learning-induced representational expansion in the primary auditory cortex (AI) directly encodes the degree of behavioral importance of a sound. Rats trained on an operant auditory conditioning task were variably motivated to the conditioned stimulus (CS) through different levels of water deprivation. Mean performance values correlated with deprivation level, validating them as a measure of the overall control and, therefore, behavioral importance of the CS. Electrophysiological mapping revealed expanded representations of the CS, compared with other frequencies in experimental subjects, but not in naive or visually trained controls that received noncontingent CS tones. Importantly, representational area showed a significant positive correlation with mean performance levels for only the CS band, with significant effects for relative area in contrast to only modest changes in absolute area. CS representational expansion was asymmetric into high-frequency zones, thus performance level also was significantly correlated with the relative anterior-posterior location of the enlarged representation. An increased representation of low frequencies, related to the acoustic spectrum of the reward delivery equipment, also was discovered in both experimental and control trained subjects, supporting the conclusion that behaviorally important sounds gain representational area. Furthermore, there was a surprising reduction in total AI area for the experimental and control groups, compared with untrained naive subjects, indicating that the functional dimensions of AI are not fixed. Overall, the findings support the encoding of acquired stimulus importance based on representational size in AI.

A central tenet in brain research is that early sensory cortex is modality specific, and, only in exceptional cases, such as deaf and blind subjects or professional musicians, is influenced by other modalities. Here we describe extensive cross-modal activation in the auditory cortex of two monkeys while they performed a demanding auditory categorization task: after a cue light was turned on, monkeys could initiate a tone sequence by touching a bar and then earn a reward by releasing the bar on occurrence of a falling frequency contour in the sequence. In their primary auditory cortex and posterior belt areas, we found many acoustically responsive neurons whose firing was synchronized to the cue light or to the touch or release of the bar. Of 315 multiunits, 45 exhibited cue light-related firing, 194 exhibited firing that was related to bar touch, and 268 exhibited firing that was related to bar release. Among 60 single units, we found one neuron with cue light-related firing, 21 with bar touch-related firing, and 36 with release-related firing. This firing disappeared at individual sites when the monkeys performed a visual detection task. Our findings corroborate and extend recent findings on cross-modal activation in the auditory cortex and suggests that the auditory cortex can be activated by visual and somatosensory stimulation and by movements. We speculate that the multimodal corepresentation in the auditory cortex has arisen from the intensive practice of the subjects with the behavioral procedure and that it facilitates the performance of audiomotor tasks in proficient subjects.

Human sound localization results primarily from the processing of binaural differences in sound level and arrival time for locations in the horizontal plane (azimuth) and of spectral shape cues generated by the head and pinnae for positions in the vertical plane (elevation). The latter mechanism incorporates two processing stages: a spectral-to-spatial mapping stage and a binaural weighting stage that determines the contribution of each ear to perceived elevation as function of sound azimuth. We demonstrated recently that binaural pinna molds virtually abolish the ability to localize sound-source elevation, but, after several weeks, subjects regained normal localization performance. It is not clear which processing stage underlies this remarkable plasticity, because the auditory system could have learned the new spectral cues separately for each ear (spatial-mapping adaptation) or for one ear only, while extending its contribution into the contralateral hemifield (binaural-weighting adaptation). To dissociate these possibilities, we applied a long-term monaural spectral perturbation in 13 subjects. Our results show that, in eight experiments, listeners learned to localize accurately with new spectral cues that differed substantially from those provided by their own ears. Interestingly, five subjects, whose spectral cues were not sufficiently perturbed, never yielded stable localization performance. Our findings indicate that the analysis of spectral cues may involve a correlation process between the sensory input and a stored spectral representation of the subject's ears and that learning acts predominantly at a spectral-to-spatial mapping level rather than at the level of binaural weighting.

Perceptual consequences of disrupted auditory nerve activity were systematically studied in 21 subjects who had been clinically diagnosed with auditory neuropathy (AN), a recently defined disorder characterized by normal outer hair cell function but disrupted auditory nerve function. Neurological and electrophysical evidence suggests that disrupted auditory nerve activity is due to desynchronized or reduced neural activity or both. Psychophysical measures showed that the disrupted neural activity has minimal effects on intensity-related perception, such as loudness discrimination, pitch discrimination at high frequencies, and sound localization using interaural level differences. In contrast, the disrupted neural activity significantly impairs timing related perception, such as pitch discrimination at low frequencies, temporal integration, gap detection, temporal modulation detection, backward and forward masking, signal detection in noise, binaural beats, and sound localization using interaural time differences. These perceptual consequences are the opposite of what is typically observed in cochlear-impaired subjects who have impaired intensity perception but relatively normal temporal processing after taking their impaired intensity perception into account. These differences in perceptual consequences between auditory neuropathy and cochlear damage suggest the use of different neural codes in auditory perception: a suboptimal spike count code for intensity processing, a synchronized spike code for temporal processing, and a duplex code for frequency processing. We also proposed two underlying physiological models based on desynchronized and reduced discharge in the auditory nerve to successfully account for the observed neurological and behavioral data. These methods and measures cannot differentiate between these two AN models, but future studies using electric stimulation of the auditory nerve via a cochlear implant might. These results not only show the unique contribution of neural synchrony to sensory perception but also provide guidance for translational research in terms of better diagnosis and management of human communication disorders.

Neurons in primary auditory cortex (A1) of cats show strong stimulus-specific adaptation (SSA). In probabilistic settings, in which one stimulus is common and another is rare, responses to common sounds adapt more strongly than responses to rare sounds. This SSA could be a correlate of auditory sensory memory at the level of single A1 neurons. Here we studied adaptation in A1 neurons, using three different probabilistic designs. We showed that SSA has several time scales concurrently, spanning many orders of magnitude, from hundreds of milliseconds to tens of seconds. Similar time scales are known for the auditory memory span of humans, as measured both psychophysically and using evoked potentials. A simple model, with linear dependence on both short-term and long-term stimulus history, provided a good fit to A1 responses. Auditory thalamus neurons did not show SSA, and their responses were poorly fitted by the same model. In addition, SSA increased the proportion of failures in the responses of A1 neurons to the adapting stimulus. Finally, SSA caused a bias in the neuronal responses to unbiased stimuli, enhancing the responses to eccentric stimuli. Therefore, we propose that a major function of SSA in A1 neurons is to encode auditory sensory memory on multiple time scales. This SSA might play a role in stream segregation and in binding of auditory objects over many time scales, a property that is crucial for processing of natural auditory scenes in cats and of speech and music in humans.

Monaurally deaf people lack the binaural acoustic difference cues in sound level and timing that are needed to encode sound location in the horizontal plane (azimuth). It has been proposed that these people therefore rely on spectral pinna cues of their normal ear to localize sounds. However, the acoustic head-shadow effect (HSE) might also serve as an azimuth cue, despite its ambiguity when absolute sound levels are unknown. Here, we assess the contribution of either cue in the monaural deaf to two-dimensional (2D) sound localization. In a localization test with randomly interleaved sound levels, we show that all monaurally deaf listeners relied heavily on the HSE, whereas binaural control listeners ignore this cue. However, some monaural listeners responded partly to actual sound-source azimuth, regardless of sound level. We show that these listeners extracted azimuth information from their pinna cues. The better monaural listeners were able to localize azimuth on the basis of spectral cues, the better their ability to also localize sound-source elevation. In a subsequent localization experiment with one fixed sound level, monaural listeners rapidly adopted a strategy on the basis of the HSE. We conclude that monaural spectral cues are not sufficient for adequate 2D sound localization under unfamiliar acoustic conditions. Thus, monaural listeners strongly rely on the ambiguous HSE, which may help them to cope with familiar acoustic environments.

The contention that normally binaural listeners can localize sound under monaural conditions has been challenged by Wightman and Kistler (J. Acoust. Soc. Am. 101:1050-1063, 1997), who found that listeners are almost completely unable to localize virtual sources of sound when sound is presented to only one ear. Wightman and Kistler's results raise the question of whether monaural spectral cues are used by listeners to localize sound under binaural conditions. We have examined the possibility that monaural spectral cues provide useful information regarding sound-source elevation and front-back hemifield when interaural time differences are available to specify sound-source lateral angle. The accuracy with which elevation and front-back hemifield could be determined was compared between a monaural condition and a binaural condition in which a wide-band signal was presented to the near ear and a version of the signal that had been lowpass-filtered at 2.5 kHz was presented to the far ear. It was found that accuracy was substantially greater in the latter condition, suggesting that information regarding sound-source lateral angle is required for monaural spectral cues to elevation and front-back hemifield to be correctly interpreted.

The term informational masking has traditionally been used to refer to elevations in signal threshold resulting from masker uncertainty. In the present study, the method of constant stimuli was used to obtain complete psychometric functions (PFs) from 44 normal-hearing listeners in conditions known to produce varying amounts of informational masking. The listener’s task was to detect a pure-tone signal in the presence of a broadband noise masker (low masker uncertainty) and in the presence of multitone maskers with frequencies and amplitudes that varied at random from one presentation to the next (high masker uncertainty). Relative to the broadband noise condition, significant reductions were observed in both the slope and the upper asymptote of the PF for multitone maskers producing large amounts of informational masking. Slope was affected more for some listeners and conditions while asymptote was affected more for others; consequently, neither parameter alone was highly predictive of individual thresholds or the amount of informational masking. Mean slopes and asymptotes varied nonmonotonically with the number of masker components in a manner similar to mean thresholds, particularly when the estimated effect of energetic masking on thresholds was subtracted out. As in past studies, the threshold data were well described by a model in which trial-by-trial judgments are based on a weighted sum of levels in dB at the output of independent auditory filters. The psychometric data, however, complicated the model’s interpretation in two ways: First, they suggested that, depending on the listener and condition, the weights can either reflect a fixed influence of masker components on each trial or the effect of occasionally mistaking a masker component for the signal from trial to trial. Second, they indicated that in either case the variance of the underlying decision variable as estimated from PF slope is not by itself great enough to account for the observed changes in informational masking.

The present study aimed to examine the effect of noise on vowel-like suprathreshold discrimination in cochlear-implant listeners. The task was to detect an increment in level at the middle harmonic (400 Hz) in the background of a seven-harmonic complex from 100 to 700 Hz in 100-Hz steps. The task was performed in the absence (control) and presence of a white noise presented over a 20-35-dB range from inaudible to loud. The present result shows that discrimination of suprathreshold harmonic stimuli was significantly enhanced, particularly at the soft signal level, with suprathreshold noise. This result suggests that tuning of the noise level is required to optimize performance of different tasks in cochlear implants.

When normal-hearing adults and children are required to detect a 1000-Hz tone in a random-frequency multitone masker, masking is often observed in excess of that predicted by traditional auditory filter models. The excess masking is called informational masking. Though individual differences in the effect are large, the amount of informational masking is typically much greater in young children than in adults [Oh et al., J. Acoust. Soc. Am. 109, 2888–2895 (2001)]. One factor that reduces informational masking in adults is spatial separation of the target tone and masker. The present study was undertaken to determine whether or not a similar effect of spatial separation is observed in children. An extreme case of spatial separation was used in which the target tone was presented to one ear and the random multitone masker to the other ear. This condition resulted in nearly complete elimination of masking in adults. In young children, however, presenting the masker to the nontarget ear typically produced only a slight decrease in overall masking and no change in informational masking. The results for children are interpreted in terms of a model that gives equal weight to the auditory filter outputs from each ear.

Masked threshold for a pure-tone signal can be substantially elevated whenever the listener is uncertain about the spectral or temporal properties of the masker, an effect referred to as auditory informational masking. Individual differences in the effect are large, with young children being most susceptible. When masker uncertainty is introduced by randomizing the frequencies of a multitone masker on each presentation, the function relating a child's pure-tone signal threshold to the number of masker components is found to be substantially elevated above that of most adults. The age effect and the individual differences among adults are not well understood, though a difference in the shapes of the masking functions suggests that different detection strategies may be involved. The present study reports results from a principal components analysis of informational masking functions obtained from 38 normal-hearing children ranging in age from 4 to 16 years and 46 normal-hearing adults ranging in age from 19 to 38 years. The premise underlying the analysis is that if different detection strategies are involved, they should add independent sources of variance to the masking functions. Hence, more than one principal component (PC) should be required to account for a substantial proportion of the variance in these functions. The results, instead, supported the operation of a single underlying strategy with all but 17% of the variance accounted for by the first PC within and across age groups. An analysis of variance on the first two PCs showed that only the first changed with age, and a cluster analysis of the masking functions showed complete separation of clusters along this PC for all but 1 listener. The results are taken to suggest that large individual differences informational masking at all ages reflect differences in the extent to which masker uncertainty adds variance to the decision variable of an otherwise optimal decision strategy.

When the source of a tone moves with respect to a listener’s ears, dichotic (or interaural) phase and amplitude modulations (PM and AM) are produced. Two experiments investigated the psychophysical characteristics of dichotic linear ramp modulations in phase and amplitude, and compared them with the psychophysics of diotic PM and AM. In experiment 1, subjects were substantially more sensitive to dichotic PM than diotic PM, but AM sensitivity was equivalent in the dichotic and diotic conditions. Thresholds for discriminating modulation direction were smaller than detection thresholds for dichotic AM, and both diotic AM and PM. Dichotic PM discrimination thresholds were similar to detection thresholds. In experiment 2, the effects of ramp duration were examined. Sensitivity to dichotic AM and PM, and diotic AM increased as duration was increased from 20 ms to 200 ms. The functions relating sensitivity to ramp duration differed across the stimuli; sensitivity to dichotic PM increased more rapidly than sensitivity to dichotic or diotic AM. This was also reflected in shorter time-constants and minimum integration times for dichotic PM detection. These findings support the hypothesis that the analysis of dichotic PM and AM rely on separate mechanisms.

Many fish rely on sounds for communication, yet the peripheral structures containing the hair cells are simple, without the morphological specializations that facilitate frequency analysis in the mammalian cochlea. Despite this, neurons in the midbrain of sound-producing fish ( Pollimyrus ) have complex receptive fields, extracting features from courtship sounds. Here we present an analysis of the initial encoding of sounds by the primary afferents and demonstrate that the representation of sound undergoes a substantial transformation as it ascends to the midbrain. Afferents were isolated as they coursed from the sacculus through the medulla. Tones (100 Hz-1.2 kHz) elicited synchronized spikes [vector strength (VS) >0.9] on each stimulus cycle [coefficient of variation (CV) <1.1], with little spike rate adaptation. Most afferents (67%) were spontaneously active and began synchronizing 10 dB below rate threshold. Rate thresholds for the most sensitive afferents (65 dB) were close to behavioral thresholds. The distribution of characteristic frequencies and best sensitivities was matched to the spectrum of sounds of this species and to its audiogram. Three clusters of afferents were identified, one including afferents that generated spike bursts and had v-shaped response areas (bursters), and two others that included entrained afferents with broad response areas (entrained types I and II). All afferents encoded the timing of clicks within click trains with time-locked spikes, and none showed selectivity for interclick intervals. Understanding the computations that yield complex receptive fields is an essential goal for auditory neuroscience, and these data on primary encoding advance this goal, allowing a comparison of inputs with feature-extracting midbrain neurons.

The interaural time difference (ITD) is a major cue to sound localization along the horizontal plane. The maximum natural ITD occurs when a sound source is positioned opposite to one ear. We examined the ability of owls and humans to detect large ITDs in sounds presented through headphones. Stimuli consisted of either broad or narrow bands of Gaussian noise, 100 ms in duration. Using headphones allowed presentation of ITDs that are greater than the maximum natural ITD. Owls were able to discriminate a sound leading to the left ear from one leading to the right ear, for ITDs that are 5 times the maximum natural delay. Neural recordings from optic-tectum neurons, however, show that best ITDs are usually well within the natural range and are never as large as ITDs that are behaviorally discriminable. A model of binaural crosscorrelation with short delay lines is shown to explain behavioral detection of large ITDs. The model uses curved trajectories of a cross-correlation pattern as the basis for detection. These trajectories represent side peaks of neural ITD-tuning curves and successfully predict localization reversals by both owls and human subjects.

Preschoolers and adults were asked to detect a 1000-Hz signal, which was masked by a multitone complex. The frequencies and amplitudes of the components in the complex varied randomly and independently on each presentation. A staircase, cued two-interval, forced-choice procedure disguised as a “listening game” was used to obtain signal thresholds in quiet and in the presence of the multitone maskers. The number of components in the masker was fixed within an experimental condition and varied from 2 to 906 across experimental conditions. Thresholds were also measured with a broadband noise masker. Eight preschool children and eight adults were tested. Although individual differences were large, among both adults and children, there was little difference between the groups in the mean amount of masking produced by the maskers with large numbers of components (400 and 906). There was also a small but significant difference between adults and children in the mean amount of masking produced by the broadband noise. The difference between the groups was much larger with smaller numbers of components. Data obtained from the adults were basically similar to that previously reported [cf. Neff and Green, Percept. Psychophys. 41, 409–415 (1987); Oh and Lutfi, J. Acoust. Soc. Am. 104, 3489–3499 (1998)]: maskers comprised of 10–40 components produced as much as 30 to 60 dB of masking in some, but not all listeners. Those same maskers produced larger amounts of masking (70–83 dB) in many of the preschool children, although, as in the adult group, individual differences were large. The component-relative-entropy (CoRE) model [Lutfi, J. Acoust. Soc. Am. 94, 748–758 (1993)] was used to describe the differences in performance between the children and adults. According to this model the average child appears to integrate information over a larger number of auditory filters than the average adult.

The relationship between sensory sensitivity and reading performance was examined to test the hypothesis that the orthographic and phonological skills engaged in visual word recognition are constrained by the ability to detect dynamic visual and auditory events. A test battery using sensory psychophysics, psychometric tests, and measures of component literacy skills was administered to 32 unselected 10-year-old primary school children. The results suggest that children's sensitivity to both dynamic auditory and visual stimuli are related to their literacy skills. Importantly, after controlling for intelligence and overall reading ability, visual motion sensitivity explained independent variance in orthographic skill but not phonological ability, and auditory FM sensitivity covaried with phonological skill but not orthographic skill. These results support the hypothesis that sensitivity at detecting dynamic stimuli influences normal children's reading skills. Vision and audition separately may affect the ability to extract orthographic and phonological information during reading.