Psychoacoustics publications

This study examines patterns of neural activity in response to single biphasic electrical pulses, presented alone or following a forward masking pulse train, delivered by a cochlear implant. Recordings were made along the tonotopic axis of the central nucleus of the inferior colliculus (ICC) in ketamine/xylazine anesthetized guinea pigs. The partial tripolar electrode configuration was used, which provided a systematic way to vary the tonotopic extent of ICC activation between monopolar (broad) and tripolar (narrow) extremes while maintaining the same peak of activation. The forward masking paradigm consisted of a 200 ms masker pulse train (1017 pulses per second) followed 10 ms later by a single-pulse probe stimulus; the current fraction of the probe was set to 0 (monopolar), 1 (tripolar), or 0.5 (hybrid), and the fraction of the masker was fixed at 0.5. Forward masking tuning profiles were derived from the amount of masking current required to just suppress the activity produced by a fixed-level probe. These profiles were sharper for more focused probe configurations, approximating the pattern of neural activity elicited by single (non-masked) pulses. The result helps to bridge the gap between previous findings in animals and recent psychophysical data.

The efficient-coding hypothesis asserts that neural and perceptual sensitivity evolved to faithfully represent biologically relevant sensory signals. Here we characterized the spectrotemporal modulation statistics of several natural sound ensembles and examined how neurons encode these statistics in the central nucleus of the inferior colliculus (CNIC) of cats. We report that modulation-tuning in the CNIC is matched to equalize the modulation power of natural sounds. Specifically, natural sounds exhibited a tradeoff between spectral and temporal modulations, which manifests as 1/ f modulation power spectrum (MPS). Neural tuning was highly overlapped with the natural sound MPS and neurons approximated proportional resolution filters where modulation bandwidths scaled with characteristic modulation frequencies, a behavior previously described in human psychoacoustics. We demonstrate that this neural scaling opposes the 1/ f scaling of natural sounds and enhances the natural sound representation by equalizing their MPS. Modulation tuning in the CNIC may thus have evolved to represent natural sound modulations in a manner consistent with efficiency principles and the resulting characteristics likely underlie perceptual resolution.

Noise-vocoded (NV) speech is often regarded as conveying phonetic information primarily through temporal-envelope cues rather than spectral cues. However, listeners may infer the formant frequencies in the vocal-tract output—a key source of phonetic detail—from across-band differences in amplitude when speech is processed through a small number of channels. The potential utility of this spectral information was assessed for NV speech created by filtering sentences into six frequency bands, and using the amplitude envelope of each band (≤30 Hz) to modulate a matched noise-band carrier (N). Bands were paired, corresponding to F1 (≈N1 + N2), F2 (≈N3 + N4) and the higher formants (F3′ ≈ N5 + N6), such that the frequency contour of each formant was implied by variations in relative amplitude between bands within the corresponding pair. Three-formant analogues (F0 = 150 Hz) of the NV stimuli were synthesized using frame-by-frame reconstruction of the frequency and amplitude of each formant. These analogues were less intelligible than the NV stimuli or analogues created using contours extracted from spectrograms of the original sentences, but more intelligible than when the frequency contours were replaced with constant (mean) values. Across-band comparisons of amplitude envelopes in NV speech can provide phonetically important information about the frequency contours of the underlying formants.

Recognition of acoustic signals may be impeded by two factors: extrinsic noise, which degrades sounds before they arrive at the receiver's ears, and intrinsic neuronal noise, which reveals itself in the trial-to-trial variability of the responses to identical sounds. Here we analyzed how these two noise sources affect the recognition of acoustic signals from potential mates in grasshoppers. By progressively corrupting the envelope of a female song, we determined the critical degradation level at which males failed to recognize a courtship call in behavioral experiments. Using the same stimuli, we recorded intracellularly from auditory neurons at three different processing levels, and quantified the corresponding changes in spike train patterns by a spike train metric, which assigns a distance between spike trains. Unexpectedly, for most neurons, intrinsic variability accounted for the main part of the metric distance between spike trains, even at the strongest degradation levels. At consecutive levels of processing, intrinsic variability increased, while the sensitivity to external noise decreased. We followed two approaches to determine critical degradation levels from spike train dissimilarities, and compared the results with the limits of signal recognition measured in behaving animals.

Whenever an acoustic scene contains a mixture of sources, listeners must segregate the mixture in order to compute source content and/or location. Some past studies have explored whether perceived location depends on which sound elements are perceived within a source. However, no direct comparisons have been made of “what” and “where” judgments for the same sound mixtures using the same listeners. The current study tested if the sound elements making up an auditory object predict that object’s perceived location. Listeners were presented with an auditory scene containing competing “target” and “captor” sources, each of which could logically contain a “promiscuous” tone complex. In separate blocks, the same listeners matched the perceived spectro-temporal content (“what”) and location (“where”) of the target. Generally, as the captor intensity decreased, the promiscuous complex contributed more to both what and where judgments of the target. However judgments did not agree either quantitatively or qualitatively. For some subjects, the promiscuous complex consistently contributed more to the spectro-temporal content of the target than to its location while for some it consistently contributed more to target location. These results show a dissociation between the perceived spectro-temporal content of an auditory object and where that object is perceived.

This study investigated the relative contributions of consonants and vowels to the perceptual intelligibility of monosyllabic consonant-vowel-consonant (CVC) words. A noise replacement paradigm presented CVCs with only consonants or only vowels preserved. Results demonstrated no difference between overall word accuracy in these conditions; however, different error patterns were observed. A significant effect of lexical difficulty was demonstrated for both types of replacement, whereas the noise level used during replacement did not influence results. The contribution of consonant and vowel transitional information present at the consonant-vowel boundary was also explored. The proportion of speech presented, regardless of the segmental condition, overwhelmingly predicted performance. Comparisons were made with previous segment replacement results using sentences [Fogerty, and Kewley-Port (2009). J. Acoust. Soc. Am. 126, 847–857]. Results demonstrated that consonants contribute to intelligibility equally in both isolated CVC words and sentences. However, vowel contributions were mediated by context, with greater contributions to intelligibility in sentence contexts. Therefore, it appears that vowels in sentences carry unique speech cues that greatly facilitate intelligibility which are not informative and/or present during isolated word contexts. Consonants appear to provide speech cues that are equally available and informative during sentence and isolated word presentations.

Speech perception requires the rapid and effortless extraction of meaningful phonetic information from a highly variable acoustic signal. A powerful example of this phenomenon is categorical speech perception, in which a continuum of acoustically varying sounds is transformed into perceptually distinct phoneme categories. We found that the neural representation of speech sounds is categorically organized in the human posterior superior temporal gyrus. Using intracranial high-density cortical surface arrays, we found that listening to synthesized speech stimuli varying in small and acoustically equal steps evoked distinct and invariant cortical population response patterns that were organized by their sensitivities to critical acoustic features. Phonetic category boundaries were similar between neurometric and psychometric functions. Although speech-sound responses were distributed, spatially discrete cortical loci were found to underlie specific phonetic discrimination. Our results provide direct evidence for acoustic-to-higher order phonetic level encoding of speech sounds in human language receptive cortex.

Objective: The purpose of this investigation was to compare the ability of young and older adults to integrate auditory and visual sentence materials under conditions of good and poor signal clarity. The principle of inverse effectiveness (PoIE), which characterizes many neuronal and behavioral phenomena related to multisensory integration, asserts that as unimodal performance declines, integration is enhanced. Thus, the PoIE predicts that both young and older adults will show enhanced integration of auditory and visual speech stimuli when these stimuli are degraded. More importantly, because older adults' unimodal speech recognition skills decline in both the auditory and visual domains, the PoIE predicts that older adults will show enhanced integration during audiovisual speech recognition relative to younger adults. This study provides a test of these predictions. Design: Fifty-three young and 53 older adults with normal hearing completed the closed-set Build-A-Sentence test and the CUNY Sentence test in a total of eight conditions; four unimodal and four audiovisual. In the unimodal conditions, stimuli were either auditory or visual and either easier or harder to perceive; the audiovisual conditions were formed from all the combinations of the unimodal signals. The hard visual signals were created by degrading video contrast, and the hard auditory signals were created by decreasing the signal to noise ratio. Scores from the unimodal and bimodal conditions were used to compute auditory enhancement and integration enhancement measures. Results: Contrary to the PoIE, neither the auditory enhancement nor integration enhancement measures increased when signal clarity in the auditory or visual channel of audiovisual speech stimuli was decreased, nor was either measure higher for older adults than for young adults. In audiovisual conditions with easy visual stimuli, the integration enhancement measure for older adults was equivalent to that for young adults. However, in conditions with hard visual stimuli, integration enhancement for older adults was significantly lower than that for young adults. Conclusions: The present findings do not support extension of the PoIE to audiovisual speech recognition. Our results are not consistent with either the prediction that integration would be enhanced under conditions of poor signal clarity or the prediction that older adults would show enhanced integration, relative to young adults. Although there is a considerable controversy with regard to the best way to measure audiovisual integration, the fact that two of the most prominent measures, auditory enhancement and integration enhancement, both yielded results inconsistent with the PoIE, strongly suggests that the integration of audiovisual speech stimuli differs in some fundamental way from the integration of other bimodal stimuli. The results also suggest that aging does not impair integration enhancement when the visual speech signal has good clarity, but may affect it when the visual speech signal has poor clarity.

The precedence effect refers to the fact that humans are able to localize sound in reverberant environments, because the auditory system assigns greater weight to the direct sound (lead) than the later-arriving sound (lag). In this study, absolute sound localization was studied for single source stimuli and for dual source lead-lag stimuli in 4–5 year old children and adults. Lead-lag delays ranged from 5–100 ms. Testing was conducted in free field, with pink noise bursts emitted from loudspeakers positioned on a horizontal arc in the frontal field. Listeners indicated how many sounds were heard and the perceived location of the first- and second-heard sounds. Results suggest that at short delays (up to 10 ms), the lead dominates sound localization strongly at both ages, and localization errors are similar to those with single-source stimuli. At longer delays errors can be large, stemming from over-integration of the lead and lag, interchanging of perceived locations of the first-heard and second-heard sounds due to temporal order confusion, and dominance of the lead over the lag. The errors are greater for children than adults. Results are discussed in the context of maturation of auditory and non-auditory factors.

Belgian Waterslager (BW) canaries have an inherited hearing loss due to missing and abnormal hair cells, but it is unclear whether the loss is congenital or developmental. We used auditory brainstem responses and scanning electron microscopy to describe the development of auditory sensitivity and hair cell abnormalities in BW and non-BW canaries. In both strains, adult ABR thresholds were higher than behavioral thresholds, but BW canaries exhibited higher thresholds than non-BW canaries across all frequencies. Immediately post-hatch, ABR thresholds and hair cell numbers were similar in both strains. Two weeks later, thresholds were significantly higher in BW canaries, and hair cell number progressively decreased as the birds aged. These data show that in BW canaries: the peripheral auditory system is functionally similar to non-BW canary from hatch to 2 weeks, ABR thresholds improve during this developmental period, actually becoming better than those of adults, but then worsen as the bird continues to age. Hair cell number and appearance is similar to non-BW canaries at hatch but progressively declines after 30 days of age. These data show that the hearing loss characteristic of BW canaries is, at least in part, developmental and is established by the time song learning begins.

Objectives: To measure sound source localization in children who have sequential bilateral cochlear implants (BICIs); to determine whether localization accuracy correlates with performance on a right-left discrimination task (i.e., spatial acuity); to determine whether there is a measurable bilateral benefit on a sound source identification task (i.e., localization accuracy) by comparing performance under bilateral and unilateral listening conditions; and to determine whether sound source localization continues to improve with longer durations of bilateral experience. Design: Two groups of children participated in this study: a group of 21 children who received BICIs in sequential procedures (5 to 14 years) and a group of 7 typically developing children with normal acoustic hearing (5 years). Testing was conducted in a large sound-treated booth with loudspeakers positioned on a horizontal arc with a radius of 1.2 m. Children participated in two experiments that assessed spatial hearing skills. Spatial hearing acuity was assessed with a discrimination task in which listeners determined whether a sound source was presented on the right or left side of center; the smallest angle at which performance on this task was reliably above chance is the minimum audible angle. Sound localization accuracy was assessed with a sound source identification task in which children identified the perceived position of the sound source from a multiloudspeaker array (7 or 15); errors are quantified using the root mean square (RMS) error. Results: Sound localization accuracy was highly variable among the children with BICIs, with RMS errors ranging from 19 to 56 degrees . Performance of the normal hearing group, with RMS errors ranging from 9 to 29 degrees was significantly better. Within the BICI group, in 11 of 21 children, RMS errors were smaller in the bilateral versus unilateral listening condition, indicating bilateral benefit. There was a significant correlation between spatial acuity and sound localization accuracy (R = 0.68, p < 0.01), suggesting that children who achieve small RMS errors tend to have the smallest minimum audible angles. Although there was large intersubject variability, testing of 11 children in the BICI group at two sequential visits revealed a subset of children who show improvement in spatial hearing skills over time. Conclusions: A subset of children who use sequential BICIs can acquire sound localization abilities, even after long intervals between activation of hearing in the first- and second-implanted ears. This suggests that children with activation of the second implant later in life may be capable of developing spatial hearing abilities. The large variability in performance among the children with BICIs suggests that maturation of sound localization abilities in children with BICIs may be dependent on various individual subject factors such as age of implantation and chronological age.

The pitch of the missing fundamental (F0) is one of the principal psychological attributes of human pitch perception. Behavioral responses to harmonic tone complexes having missing F0s were measured in chinchillas using operant conditioning and stimulus generalization. Animals were trained to discriminate between tone complexes having a 500-Hz F0 and a 125-Hz F0. When animals were tested with tone complexes having the same F0s, but where the F0s were missing, responses were similar to those obtained when the F0s were present, suggesting that missing F0 sounds were perceptually equivalent to F0 present sounds. Behavioral responses to F0 present and missing F0 stimuli were similar in the presence of low-pass masking noise, suggesting that the perception was not due to the reinsertion of the F0 through cochlear nonlinearities. Gradients in behavioral responses were observed when the F0s of test complexes were systematically varied, suggesting the existence of a psychological dimension related to F0. Behavioral responses were related to the F0 rather than to spectral differences among test stimuli when the F0 and spectrum were varied independently. The results indicate that chinchillas possess a pitch-like perception of the missing F0 that is unlikely to arise from cochlear distortion products.

Vision and audition provide spatial information about the environment to guide natural behavior. Because the eyes move in the head while the ears remain head-fixed, input conveying eye position in the head is required to maintain audiovisual congruence. Human perception of auditory space was previously shown to shift with changes in eye position, regardless of the target's frequency content and spatial cues underlying horizontal and vertical localization. In this study, we examined whether this interaction is altered by advancing age. Head-restrained young (18-44 yo), middle-aged (45-64 yo), and elderly (65-81 yo) human subjects localized noise bursts under conditions of transient and sustained ocular deflection. All three age groups demonstrated a time-dependent shift of auditory space in the direction of eye position. Moreover, this adaptation showed a clear decline with advancing age, but only for peripheral auditory space (beyond ±10° from midline). Alternatively, adaptation in the periphery may occur, but is more sluggish than in the central field and therefore not fully observed in this experiment. The age-dependent effect cannot be readily explained by senescent peripheral hearing loss, suggesting a change in central processing of auditory space in relation to the control of gaze.

Segregation of concurrent sounds in complex acoustic environments is a fundamental feature of auditory scene analysis. A powerful cue used by the auditory system to segregate concurrent sounds, such as speakers' voices at a cocktail party, is inharmonicity. This can be demonstrated when a component of a harmonic complex tone is perceived as a separate tone “popping out” from the complex as a whole when it is sufficiently mistuned from its harmonic value. The neural bases of perceptual “pop out” of mistuned harmonics are unclear. We recorded multiunit activity from primary auditory cortex (A1) of behaving monkeys elicited by harmonic complex tones that were either “in tune” or that contained a mistuned third harmonic set at the best frequency of the neural populations. Responses to mistuned sounds were enhanced relative to responses to “in-tune” sounds, thus correlating with the enhanced perceptual salience of the mistuned component. Consistent with human psychophysics of “pop out,” response enhancements increased with the degree of mistuning, were maximal for neural populations tuned to the frequency of the mistuned component, and were not observed under comparable stimulus conditions that do not elicit perceptual “pop out.” Mistuning was also associated with changes in neuronal temporal response patterns phase locked to “beats” in the stimuli. Intracortical auditory evoked potentials paralleled noninvasive neurophysiological correlates of perceptual “pop out” in humans, further augmenting the translational relevance of the results. Findings suggest two complementary neural mechanisms for “pop out,” based on the detection of local differences in activation level or coherence of temporal response patterns across A1.

In this study, combinations of interaural time differences (ITD) and interaural level differences (ILD) were applied to trains of 4000 Hz Gabor clicks (Gaussian-filtered impulses) and presented to listeners over headphones. ITD/ILD equivalence functions, or "trading ratios" (TR) were estimated using two different procedures: a "closed-loop" procedure in which subjects adjusted (via head-turn) the ILD of a target click train to counteract the effects of an imposed ITD, and an "open-loop" procedure in which subjects indicated (also via head-turn) the lateral position of click trains containing independent combinations of ITD and ILD. For both tasks, TR values increasingly favored ILD over ITD as inter-click interval (ICI) decreased from 10 to 2 ms. Subsequent analysis confirmed that this change reflected a loss of sensitivity to envelope ITD at short ICI rather than a gain in sensitivity to ILD, consistent with prior studies demonstrating rate-limited processing of ongoing envelope ITD. Significant intersubject differences in the data included two subjects whose TR values obtained under both procedures were consistently lower (greater influence of ITD) than other subjects', and did not vary with ICI. Such differences suggest that multiple mechanisms of ITD/ILD combination may be utilized to varying degrees by individual listeners. By at least one of those mechanisms, ITD sensitivity (but not ILD sensitivity) is limited to low modulation rates.

Songbirds, which, like humans, learn complex vocalizations, provide an excellent model for the study of acoustic pattern recognition. Here we examined the role of three basic acoustic parameters in an ethologically relevant categorization task. Female zebra finches were first trained to classify songs as belonging to one of two males and then asked whether they could generalize this knowledge to songs systematically altered with respect to frequency, timing, or intensity. Birds' performance on song categorization fell off rapidly when songs were altered in frequency or intensity, but they generalized well to songs that were changed in duration by >25%. Birds were not deaf to timing changes, however; they detected these tempo alterations when asked to discriminate between the same song played back at two different speeds. In addition, when birds were retrained with songs at many intensities, they could correctly categorize songs over a wide range of volumes. Thus although they can detect all these cues, birds attend less to tempo than to frequency or intensity cues during song categorization. These results are unexpected for several reasons: zebra finches normally encounter a wide range of song volumes but most failed to generalize across volumes in this task; males produce only slight variations in tempo, but females generalized widely over changes in song duration; and all three acoustic parameters are critical for auditory neurons. Thus behavioral data place surprising constraints on the relationship between previous experience, behavioral task, neural responses, and perception. We discuss implications for models of auditory pattern recognition.

The generalization of learning from trained to untrained conditions is of great potential value because it markedly increases the efficacy of practice. In principle, generalization and the learning itself could arise from either the same or distinct neural changes. Here, we assessed these two possibilities in the realm of human perceptual learning by comparing the time course of improvement on a trained condition (learning) to that on an untrained condition (generalization) for an auditory temporal-interval discrimination task. While significant improvement on the trained condition occurred within 2 d, generalization to the untrained condition lagged behind, only emerging after 4 d. The different time courses for learning and generalization suggest that these two types of perceptual improvement can arise from at least partially distinct neural changes. The notably longer time course for generalization than learning demonstrates that increasing the duration of training can be an effective means to increase the number of conditions to which learning generalizes on perceptual tasks.

Young normal-hearing (YNH) and older normal-hearing (ONH) listeners identified vowels in naturally produced /bVb/ syllables and in modified syllables that consisted of variable portions of the vowel edges (silent-center [SC] stimuli) or vowel center (center-only [CO] stimuli). Listeners achieved high levels of performance for all but the shortest stimuli, indicating that they were able to access vowel cues throughout the syllable. ONH listeners performed similarly to YNH listeners for most stimuli, but performed more poorly for the shortest CO stimuli. SC and CO stimuli were equally effective in supporting vowel identification except when acoustic information was limited to 20 ms.

Adult rats were trained to detect the occurrence of a two-element sound sequence in a background of nine other nontarget sound pairs. Training resulted in a modest, enduring, static expansion of the cortical areas of representation of both target stimulus sounds. More importantly, once the initial stimulus A in the target A-B sequence was presented, the cortical “map” changed dynamically, specifically to exaggerate further the representation of the “anticipated” stimulus B. If B occurred, it was represented over a larger cortical area by more strongly excited, more coordinated, and more selectively responding neurons. This biasing peaked at the expected time of B onset with respect to A onset. No dynamic biasing of responses was recorded for any sound presented in a nontarget pair. Responses to nontarget frequencies flanking the representation of B were reduced in area and in response strength only after the presentation of A at the expected time of B onset. This study shows that cortical areas are not representationally static but, to the contrary, can be biased moment by moment in time as a function of behavioral context.

Objective: To further examine the reliability of categorical loudness scaling (CLS) for individual loudness categories and for the slope of the CLS functions. And, to evaluate the relationship between CLS and audiometric threshold. Design: CLS functions were obtained in 74 subjects, 58 with hearing loss and 16 with normal hearing. CLS functions were measured at three frequencies (1, 2, and 4 kHz) in two separate sessions separated by as little as 1 wk and as much as 6 mo. Reliability of mean and median levels within each loudness category was assessed using SDs and correlation coefficients. Lines were fit to the CLS functions, and slopes of the lines were used to assess reliability and the relation between CLS and audiometric threshold. Results: Similar reliability for CLS measurements was observed in both normal-hearing and hearing-impaired subjects at all frequencies. Across both groups of subjects, correlations describing the reliability of mean stimulus level within category exceeded 0.92 at all frequencies. In addition, SDs of the mean stimulus-level difference between visits ranged from 6.6 to 7.8 dB, depending on frequency. The correlation between the slope of a straight line fitted to the entire CLS function and audiometric threshold collapsed across frequencies was 0.72. Two line segments were then fit to the CLS function: one segment was fit to the soft portion of the CLS function (categorical units < or =20) and the other segment to the loud portion (categorical unit >20). Slopes of the line fit to the entire CLS function and of the line fit to the soft portion of the CLS function were both reliable across sessions. The slope of the line fit to the soft portion increased as audiometric threshold increased, with the correlations greater than 0.86 at all frequencies. No relationship was observed between slope of the line fit to the loud portion of the CLS function and audiometric threshold. Iso-loudness contours were constructed from the CLS data and used to determine the gain that would be needed to produce "normal" loudness percepts for hearing-impaired individuals. Conclusions: Within-subject CLS measurements were reliable across sessions both for individual loudness categories and for slope of the CLS functions. In addition, the slope of the low-level portion of the CLS function varied in a predictable manner with audiometric threshold, with slope increasing as audiometric threshold increased. Finally, gain as a function of input level needed to provide loudness percepts for individuals with hearing loss equal to the loudness percepts of normal-hearing individuals can be estimated from audiometric threshold. This finding supports the assumption that audiometric threshold and response growth (loudness) are both determined by the same underlying cochlear mechanisms.