Psychoacoustics publications

This project focused on the individual differences underlying observed variability in temporal processing among older listeners. Four measures of vowel temporal-order identification were completed by young (N=35; 18–31 years) and older (N=151; 60–88 years) listeners. Experiments used forced-choice, constant-stimuli methods to determine the smallest stimulus onset asynchrony (SOA) between brief (40 or 70 ms) vowels that enabled identification of a stimulus sequence. Four words (pit, pet, pot, and put) spoken by a male talker were processed to serve as vowel stimuli. All listeners identified the vowels in isolation with better than 90% accuracy. Vowel temporal-order tasks included the following: (1) monaural two-item identification, (2) monaural four-item identification, (3) dichotic two-item vowel identification, and (4) dichotic two-item ear identification. Results indicated that older listeners had more variability and performed poorer than young listeners on vowel-identification tasks, although a large overlap in distributions was observed. Both age groups performed similarly on the dichotic ear-identification task. For both groups, the monaural four-item and dichotic two-item tasks were significantly harder than the monaural two-item task. Older listeners’ SOA thresholds improved with additional stimulus exposure and shorter dichotic stimulus durations. Individual differences of temporal-order performance among the older listeners demonstrated the influence of cognitive measures, but not audibility or age.

The ability to make intensity judgments for sequential stimuli was examined with an intensity-discrimination task involving three 50-ms noise bursts with non-overlapping frequency ranges. Targets (single bursts) presented in three-burst sequences were required to be as much as 5 dB more intense than targets presented as single bursts in isolation, especially for the later targets. Randomizing target position in the sequence did not reliably reduce performance, nor were thresholds for younger and older listeners reliably different. These increases in increment detection threshold are indications of a specific intensity-processing deficit for stimuli occurring later in a sequence.

Experiments to study voice quality have typically used rating scales or direct magnitude estimation to obtain listener judgments. Unfortunately, the data obtained using these tasks are context dependent, which makes it difficult to compare perceptual judgments of voice quality across experiments. The present experiment describes a simple matching task to quantify voice quality. The data obtained through this task were compared to perceptual judgments obtained using rating scale and direct magnitude estimation tasks to determine whether the three tasks provide equivalent perceptual distances across stimuli. Ten synthetic vowel continua that varied in terms of their aspiration noise were evaluated for breathiness using each of the three tasks. Linear and nonlinear regressions were used to compare the perceptual distances between stimuli obtained through each technique. Results show that the perceptual distances estimated from matching and direct magnitude estimation task are similar, but both differ from the rating scale task, suggesting that the matching task provides perceptual distances with ratio-level measurement properties. The matching task is advantageous for measurement of vocal quality because it provides reliable measurement with ratio-level scale properties. It allows the use of a fixed reference signal for all comparisons, thus allowing researchers to directly compare findings across different experiments.

The possibility that “dead regions” or “spectral holes” can account for some differences in performance between bilateral cochlear implant (CI) users and normal-hearing listeners was explored. Using a 20-band noise-excited vocoder to simulate CI processing, this study examined effects of spectral holes on speech reception thresholds (SRTs) and spatial release from masking (SRM) in difficult listening conditions. Prior to processing, stimuli were convolved through head-related transfer-functions to provide listeners with free-field directional cues. Processed stimuli were presented over headphones under binaural or monaural (right ear) conditions. Using Greenwood’s [(1990). J. Acoust. Soc. Am. 87, 2592–2605] frequency-position function and assuming a cochlear length of 35mm, spectral holes were created for variable sizes (6 and 10mm) and locations (base, middle, and apex). Results show that middle-frequency spectral holes were the most disruptive to SRTs, whereas high-frequency spectral holes were the most disruptive to SRM. Spectral holes generally reduced binaural advantages in difficult listening conditions. These results suggest the importance of measuring dead regions in CI users. It is possible that customized programming for bilateral CI processors based on knowledge about dead regions can enhance performance in adverse listening situations.

The lateral superior olive (LSO) is one of the earliest sites in the auditory pathway involved in processing acoustical cues to sound location. LSO neurons encode the interaural level difference (ILD) cue to azimuthal location. Here we investigated the effect of variations in the overall stimulus levels of sounds at the two ears on the sensitivity of LSO neurons to small differences in ILDs of pure tones. The neuronal firing rate versus ILD functions were found to depend greatly on the overall stimulus level, typically shifting along the ILD axis toward the excitatory ear and attaining greater maximal firing rates as stimulus level increased. Seventy-five percent of neurons showed significant shifts with changes in overall sound level. The range of ILDs corresponding to best neural acuity for ILDs shifted accordingly. In a simulation using the empirical data, when the overall stimulus level was randomly changed from one trial to the next, the neural discrimination thresholds for ILD, or ILD acuities, were worsened by 50–60% across the population of neurons relative to fixed stimulus levels whether ILD acuity was measured at the azimuthal midline or the ILD pedestal producing the best acuity. The impairment in ILD discrimination was attributed to the increased neural response variance imparted by varying the stimulus level. These results contrast to those observed in psychophysical studies where ILD discrimination thresholds under similar experimental conditions are invariant to overall changes in stimulus level. A simple computational model that incorporated the antagonistic inputs of bilateral LSO nuclei as well as the dorsal nuclei of the lateral lemniscus to the inferior colliculus produced a more robust encoding of ILD even in the setting of roving stimulus level. Testable predictions of this model and comparison to other computational models addressing stimulus invariance were considered.

The most important acoustic cues available to the brain for sound localization are produced by the interaction of sound with the animal's head and external ears. As a first step in understanding the relation between these cues and their neural representation in a vocal new-world primate, we measured head-related transfer functions (HRTFs) across frequency for a wide range of sound locations in three anesthetized marmoset monkeys. The HRTF magnitude spectrum has a broad resonance peak at 6-12 kHz that coincides with the frequency range of the major call types of this species. A prominent first spectral notch (FN) in the HRTF magnitude above this resonance was observed at most source locations. The center frequency of the FN increased monotonically from approximately 12 to 26 kHz with increases in elevation in the lateral field. In the frontal field FN frequency changed in a less orderly fashion with source position. From the HRTFs we derived interaural time (ITDs) and level differences (ILDs). ITDs and ILDs (below 12 kHz) varied as a function of azimuth between +/-250 micros and +/-20dB, respectively. A reflexive orienting behavioral paradigm was used to confirm that marmosets can orient to sound sources.

Humans are much better in relative than in absolute judgments. This common assertion is based on findings that discrimination thresholds are much lower when measured with methods that allow interstimuli comparisons than when measured with methods that require classification of one stimulus at a time and are hence sensitive to memory load. We now challenged this notion by measuring discrimination thresholds and evoked potentials while listeners performed a two-tone frequency discrimination task. We tested various protocols that differed in the pattern of cross-trial tone repetition. We found that best performance was achieved only when listeners effectively used cross-trial repetition to avoid interstimulus comparisons with the repeated reference tone. Instead, they classified one tone, the nonreference tone, as either high or low by comparing it with a recently formed internal reference. Listeners were not aware of the switch from interstimulus comparison to classification. Its successful use was revealed by the conjunction of improved behavioral performance and an event-related potential component (P3), indicating an implicit perceptual decision, which followed the nonreference tone in each trial. Interestingly, tone repetition itself did not suffice for the switch, implying that the bottleneck to discrimination does not reside at the lower, sensory stage. Rather, the temporal consistency of repetition was important, suggesting the involvement of higher-level mechanisms with longer time constants. These findings suggest that classification is based on more automatic and accurate mechanisms than interstimulus comparisons and that the ability to effectively use them depends on a dynamic interplay between higher- and lower-level cortical mechanisms.

Underwater vibratory sources simultaneously present acoustic and hydrodynamic disturbances. Because vibratory dipole sources are poor sonic projectors, most researchers have assumed that such sources are of greatest relevance to the lateral line system. Both hydroacoustic principles and empirical studies have shown that dipole sources are also a potent stimulus to the inner ear. Responses to vibratory sources in mottled sculpin (Cottus bairdi) were assessed using unconditioned orienting, differential and nondifferential conditioning. Orienting responses are dominated by lateral line inputs and eliminated by lateral line inactivation. Simple conditioning depends on inputs from other systems and was not affected by lateral line inactivation. Differential conditioning alters behavioral control, and sculpin could be conditioned to ignore substrate-borne vibrations and respond only to hydroacoustic stimulation of the ear. The lateral line and inner ear of mottled sculpin do not necessarily exhibit range fractionation, as both systems operate over a similar distance (within 1.5 body lengths) and respond to many of the same sources. Vibratory dipole sources generate compound stimuli that simultaneously activate multiple octavolateralis systems, and sculpin make use of the channels differentially under different behavioral tasks.

Cochlear implants stimulate the auditory nerve with amplitude-modulated (AM) electric pulse trains. Pulse rates >2,000 pulses per second (pps) have been hypothesized to enhance transmission of temporal information. Recent studies, however, have shown that higher pulse rates impair phase locking to sinusoidal AM in the auditory cortex and impair perceptual modulation detection. Here, we investigated the effects of high pulse rates on the temporal acuity of transmission of pulse trains to the auditory cortex. In anesthetized guinea pigs, signal-detection analysis was used to measure the thresholds for detection of gaps in pulse trains at rates of 254, 1,017, and 4,069 pps and in acoustic noise. Gap-detection thresholds decreased by an order of magnitude with increases in pulse rate from 254 to 4,069 pps. Such a pulse-rate dependence would likely influence speech reception through clinical speech processors. To elucidate the neural mechanisms of gap detection, we measured recovery from forward masking after a 196.6-ms pulse train. Recovery from masking was faster at higher carrier pulse rates and masking increased linearly with current level. We fit the data with a dual-exponential recovery function, consistent with a peripheral and a more central process. High-rate pulse trains evoked less central masking, possibly due to adaptation of the response in the auditory nerve. Neither gap detection nor forward masking varied with cortical depth, indicating that these processes are likely subcortical. These results indicate that gap detection and modulation detection are mediated by two separate neural mechanisms.

Traditionally time perception has been considered the product of a central, generic, cognitive mechanism. However, evidence is emerging for a distributive system with modality-specific sensory components (Morrone et al., 2005; Johnston et al., 2006). Here we show that fast contrast adaptation, which can be observed in the retina, induces a change in apparent duration. The perceived duration of a subsecond interval containing a 50% luminance contrast drifting pattern is compressed when it follows a high (90%) as compared to a low (10%) contrast interval. The duration effect cannot be attributed to changes in latency at onset relative to offset, can be dissociated from the effect of contrast context on apparent speed or apparent contrast per se and it occurs in a retinocentric frame of reference. The temporal compression is limited to high drift temporal frequencies (≥10 Hz) and is not observed for equiluminant chromatic stimuli. This pattern of results indicates a major role for the magnocellular pathway in the neural encoding and representation of visual time.

A form of processed speech is described that is highly discriminable in a closed-set identification format. The processing renders speech into a set of sinusoidal pulses played synchronously across frequency. The processing and results from several experiments are described. The number and width of frequency analysis channels and tone-pulse duration were variables. In one condition, various proportions of the tones were randomly removed. The processed speech was remarkably resilient to these manipulations. This type of speech may be useful for examining multitalker listening situations in which a high degree of stimulus control is required.

Previous studies have shown that residual acoustic hearing supplements cochlear implant function to improve speech recognition in noise as well as perception of music. The current study had two primary objectives. First, we sought to determine how cochlear implantation and electrical stimulation over a time period of 14 to 21 months influence cochlear structures such as hair cells and spiral ganglion neurons. Second, we sought to investigate whether the structures that provide acoustic hearing also affect the perception of electrical stimulation. We compared psychophysical responses to cochlear implant stimulation in two groups of adult guinea pigs. Group I (11 animals) received a cochlear implant in a previously untreated ear, while group II (ten animals) received a cochlear implant in an ear that had been previously infused with neomycin to destroy hearing. Psychophysical thresholds were measured in response to pulse-train and sinusoidal stimuli. Histological analysis of all group I animals and a subset of group II animals was performed. Nine of the 11 group I animals showed survival of the organ of Corti and spiral ganglion neurons adjacent to the electrode array. All group I animals showed survival of these elements in regions apical to the electrode array. Group II animals that were examined histologically showed complete loss of the organ of Corti in regions adjacent and apical to the electrode array and severe spiral ganglion neuron loss, consistent with previous reports for neomycin-treated ears. Behaviorally, group II animals had significantly lower thresholds than group I animals in response to 100 Hz sinusoidal stimuli. However, group I animals had significantly lower thresholds than group II animals in response to pulse-train stimuli (0.02 ms/phase; 156 to 5,000 pps). Additionally, the two groups showed distinct threshold versus pulse rate functions. We hypothesize that the differences in detection thresholds between groups are caused by the electrical activation of the hair cells in group I animals and/or differences between groups in the condition of the spiral ganglion neurons.

Background: When sound arrives at the eardrum it has already been filtered by the body, head, and outer ear. This process is mathematically described by the head-related transfer functions (HRTFs), which are characteristic for the spatial position of a sound source and for the individual ear. HRTFs in the barn owl (Tyto alba) are also shaped by the facial ruff, a specialization that alters interaural time differences (ITD), interaural intensity differences (ILD), and the frequency spectrum of the incoming sound to improve sound localization. Here we created novel stimuli to simulate the removal of the barn owl's ruff in a virtual acoustic environment, thus creating a situation similar to passive listening in other animals, and used these stimuli in behavioral tests. Methodology/principal findings: HRTFs were recorded from an owl before and after removal of the ruff feathers. Normal and ruff-removed conditions were created by filtering broadband noise with the HRTFs. Under normal virtual conditions, no differences in azimuthal head-turning behavior between individualized and non-individualized HRTFs were observed. The owls were able to respond differently to stimuli from the back than to stimuli from the front having the same ITD. By contrast, such a discrimination was not possible after the virtual removal of the ruff. Elevational head-turn angles were (slightly) smaller with non-individualized than with individualized HRTFs. The removal of the ruff resulted in a large decrease in elevational head-turning amplitudes. Conclusions/significance: The facial ruff a) improves azimuthal sound localization by increasing the ITD range and b) improves elevational sound localization in the frontal field by introducing a shift of iso-ILD lines out of the midsagittal plane, which causes ILDs to increase with increasing stimulus elevation. The changes at the behavioral level could be related to the changes in the binaural physical parameters that occurred after the virtual removal of the ruff. These data provide new insights into the function of external hearing structures and open up the possibility to apply the results on autonomous agents, creation of virtual auditory environments for humans, or in hearing aids.

Budgerigars and zebra finches were tested, using operant conditioning techniques, on their ability to identify a zebra finch song in the presence of a background masker emitted from either the same or a different location as the signal. Identification thresholds were obtained for three masker types differing in their spectrotemporal characteristics (noise, modulated noise, and a song chorus). Both bird species exhibited similar amounts of spatial unmasking across the three masker types. The amount of unmasking was greater when the masker was played continuously compared to when the target and masker were presented simultaneously. These results suggest that spatial factors are important for birds in the identification of natural signals in noisy environments.

Some past studies suggest that when sound elements are heard as one object, the spatial cues in the component elements are integrated to determine perceived location, and that this integration is reduced when the elements are perceived in separate objects. The current study explored how object localization depends on the spatial, spectral, and temporal configurations of sound elements in an auditory scene. Localization results are interpreted in light of results from a series of previous experiments studying perceptual grouping of the same stimuli, e.g., Shinn-Cunningham et al. [Proc. Natl. Acad. Sci. U.S.A. 104, 12223–12227 (2007)]. The current results suggest that the integration (pulling) of spatial information across spectrally interleaved elements is obligatory when these elements are simultaneous, even though past results show that these simultaneous sound elements are not grouped strongly into a single perceptual object. In contrast, perceptually distinct objects repel (push) each other spatially with a strength that decreases as the temporal separation between competing objects increases. These results show that the perceived location of an attended object is not easily predicted by knowledge of how sound elements contribute to the perceived spectro-temporal content of that object.

Understanding the mechanistic substrates of neural computations that lead to behavior remains a fundamental problem in neuroscience. In particular, the contributions of intrinsic neural properties such as burst firing and dendritic morphology to the processing of behaviorally relevant sensory input have received much interest recently. Pyramidal cells within the electrosensory lateral line lobe of weakly electric fish display an intrinsic bursting mechanism that relies on somato-dendritic interactions when recorded in vitro: backpropagating somatic action potentials trigger dendritic action potentials that lead to a depolarizing afterpotential (DAP) at the soma. We recorded intracellularly from these neurons in vivo and found firing patterns that were quite different from those seen in vitro: we found no evidence for DAPs as each somatic action potential was followed by a pronounced afterhyperpolarization (AHP). Calcium chelators injected in vivo reduced the AHP, thereby unmasking the DAP and inducing in vitro-like bursting in pyramidal cells. These bursting dynamics significantly reduced the cell's ability to encode the detailed time course of sensory input. We performed additional in vivo pharmacological manipulations and mathematical modeling to show that calcium influx through N-methyl-d-aspartate (NMDA) receptors activate dendritic small conductance (SK) calcium-activated potassium channels, which causes an AHP that counteracts the DAP and leads to early termination of the burst. Our results show that ion channels located in dendrites can have a profound influence on the processing of sensory input by neurons in vivo through the modulation of an intrinsic bursting mechanism.

Deafness is the most common sensory disorder in humans and the aetiology of genetic deafness is complex. Mouse mutants have been crucial in identifying genes involved in hearing. However, many deafness genes remain unidentified. Using N ‐ethyl N −nitrosourea (ENU) mutagenesis to generate new mouse models of deafness, we identified a novel semi‐dominant mouse mutant, Cloth‐ears ( Clth ). Cloth‐ears mice show reduced acoustic startle response and mild hearing loss from ∼30 days old. Auditory‐evoked brainstem response (ABR) and distortion product otoacoustic emission (DPOAE) analyses indicate that the peripheral neural auditory pathway is impaired in Cloth‐ears mice, but that cochlear function is normal. In addition, both Clth/Clth and Clth/+ mice display paroxysmal tremor episodes with behavioural arrest. Clth/Clth mice also show a milder continuous tremor during movement and rest. Longitudinal phenotypic analysis showed that Clth/+ and Clth/Clth mice also have complex defects in behaviour, growth, neurological and motor function. Positional cloning of Cloth‐ears identified a point mutation in the neuronal voltage‐gated sodium channel α‐subunit gene, Scn8a , causing an aspartic acid to valine (D981V) change six amino acids downstream of the sixth transmembrane segment of the second domain (D2S6). Complementation testing with a known Scn8a mouse mutant confirmed that this mutation is responsible for the Cloth‐ears phenotype. Our findings suggest a novel role for Scn8a in peripheral neural hearing loss and paroxysmal motor dysfunction.

Objectives: The objective of this study was to evaluate the efficacy of a word-based auditory-training procedure for use with older adults who have impaired hearing. The emphasis during training and assessment is placed on words with a high frequency of occurrence in American English. Design: In this study, a repeated-measures group design was used with each of the two groups of participants to evaluate the effects of the word-based training regimen. One group comprised 20 young adults with normal hearing and the other consisted of 16 older adults with impaired hearing. The group of young adults was not included for the purpose of between-group comparisons. Rather, it was included to demonstrate the efficacy of the training regimen, should efficacy fail to be demonstrated in the group of older adults, and to estimate the magnitude of the benefits that could be achieved in younger listeners. Results: Significant improvements were observed in the group means for each of five measures of post-training assessment. Pretraining and post-training performance assessments were all based on the open-set recognition of speech in a fluctuating speech-like background noise. Assessment measures ranged from recognition of trained words and phrases produced by talkers heard during training to the recognition of untrained sentences produced by a talker not encountered during training. In addition to these group data, analysis of individual data via 95% critical differences for each assessment measure revealed that 75 to 80% of the older adults demonstrated significant improvements on most or all of the post-training measures. Conclusions: The word-based auditory-training program examined here, one based on words having a high frequency of occurrence in American English, has been demonstrated to be efficacious in older adults with impaired hearing. Training on frequent words and frequent phrases generalized to sentences constructed from frequently occurring words whether spoken by talkers heard during training or by a novel talker.

Subject responses were measured for individual narrow-band reproducible stimuli in a low-frequency tone-in-noise detection task. Both N0S0 and N0Sπ conditions were examined. The goal of the experiment was to determine the relative importance of envelope and fine-structure cues. Therefore, chimeric stimuli were generated by recombining envelopes and fine structures from different reproducible stimuli. Detection judgments for noise-alone or tone-plus-noise stimuli that had common envelopes but different fine structures or common fine structures but different envelopes were compared. The results showed similar patterns of responses to stimuli that shared envelopes, indicating the importance of envelope cues; however, fine-structure cues were also shown to be important. The relative weight assigned to envelope and fine-structure cues varied across subjects and across interaural conditions. The results also indicated that envelope and fine-structure information are not processed independently. Implications for monaural and binaural models of masking are discussed.

This study examined the effects of increased processing load on the closed‐set speech‐identification performance of young and older adults in a one‐talker background. Since the older adults had impaired hearing, speech‐identification performance was measured for spectrally shaped stimuli comparable to those experienced when wearing well‐fit hearing aids. There were three groups of listeners: (1) 19 older adults with high‐frequency sensorineural hearing loss; (2) 10 young adults with normal hearing who were assessed with the same spectrally shaped stimuli as the older adults; (3) 9 young adults with normal hearing who were assessed without spectral shaping and at a poorer target‐to‐competition ratio in an effort to equate overall performance to that of the older adults. In addition to this group factor, there were three within‐participant repeated‐measures independent variables designed to increase the demands on processing for the target and competing speech stimuli. These were: (1) competition meaningfulness (played in forward or reverse direction); (2) gender match between target and competing talkers (same or different gender); and (3) talker uncertainty (either the same target/competition talker pair or one of many such pairs on each trial). These three repeated‐measures independent variables were examined in a 2 × 2 × 2 factorial design. They showed roughly independent and additive effects on speech‐identification such that combinations of these variables decreased performance cumulatively. Older adults performed worse than young adults across the board, but also showed diminished relative improvement as the processing load was decreased. Individual differences in performance among the older adults were also examined.