Psychoacoustics publications

A key challenge in neuroscience is understanding how sensory stimuli give rise to perception, especially when the process is supported by neural activity from an extended network of brain areas. Perception is inherently subjective, so interrogating its neural signatures requires, ideally, a combination of three factors: (1) behavioral tasks that separate stimulus-driven activity from perception per se; (2) human subjects who self-report their percepts while performing those tasks; and (3) concurrent neural recordings acquired at high spatial and temporal resolution. In this study, we analyzed human electrocorticographic recordings obtained during an auditory task which supported mutually exclusive perceptual interpretations. Eight neurosurgical patients (5 male; 3 female) listened to sequences of repeated triplets where tones were separated in frequency by several semitones. Subjects reported spontaneous alternations between two auditory perceptual states, 1-stream and 2-stream, by pressing a button. We compared averaged auditory evoked potentials (AEPs) associated with 1-stream and 2-stream percepts and identified significant differences between them in primary and nonprimary auditory cortex, surrounding auditory-related temporoparietal cortex, and frontal areas. We developed classifiers to identify spatial maps of percept-related differences in the AEP, corroborating findings from statistical analysis. We used one-dimensional embedding spaces to perform the group-level analysis. Our data illustrate exemplar high temporal resolution AEP waveforms in auditory core region; explain inconsistencies in perceptual effects within auditory cortex, reported across noninvasive studies of streaming of triplets; show percept-related changes in frontoparietal areas previously highlighted by studies that focused on perceptual transitions; and demonstrate that auditory cortex encodes maintenance of percepts and switches between them.SIGNIFICANCE STATEMENT The human brain has the remarkable ability to discern complex and ambiguous stimuli from the external world by parsing mixed inputs into interpretable segments. However, one's perception can deviate from objective reality. But how do perceptual discrepancies occur? What are their anatomical substrates? To address these questions, we performed intracranial recordings in neurosurgical patients as they reported their perception of sounds associated with two mutually exclusive interpretations. We identified signatures of subjective percepts as distinct from sound-driven brain activity in core and non-core auditory cortex and frontoparietal cortex. These findings were compared with previous studies of auditory bistable perception and suggested that perceptual transitions and maintenance of perceptual states were supported by common neural substrates.

The extent to which the primary auditory cortex (A1) participates in instructing animal behavior remains debated. Although multiple studies have shown A1 activity to correlate with animals' perceptual judgments (Jaramillo and Zador, 2011; Bizley et al., 2013; Rodgers and DeWeese, 2014), others have found no relationship between A1 responses and reported auditory percepts (Lemus et al., 2009; Dong et al., 2011). To address this ambiguity, we performed chronic recordings of evoked local field potentials (eLFPs) in A1 of head-fixed female rats performing a two-alternative forced-choice auditory discrimination task. Rats were presented with two interleaved sequences of pure tones from opposite sides and had to indicate the side from which the higher-frequency target stimulus was played. Animal performance closely correlated (rrm = 0.68) with the difference between the target and distractor eLFP responses: the more the target response exceeded the distractor response, the better the animals were at identifying the side of the target frequency. Reducing the evoked response of either frequency through stimulus-specific adaptation affected performance in the expected way: target localization accuracy was degraded when the target frequency was adapted and improved when the distractor frequency was adapted. Target frequency eLFPs were stronger on hit trials than on error trials. Our results suggest that the degree to which one stimulus stands out over others within A1 activity may determine its perceptual saliency for the animals and accordingly bias their behavioral choices.SIGNIFICANCE STATEMENT The brain must continuously calibrate the saliency of sensory percepts against their relevance to the current behavioral goal. The inability to ignore irrelevant distractors characterizes a spectrum of human attentional disorders. Meanwhile, the connection between the neural underpinnings of stimulus saliency and sensory decisions remains elusive. Here, we record local field potentials in the primary auditory cortex of rats engaged in auditory discrimination to investigate how the cortical representation of target and distractor stimuli impacts behavior. We find that the amplitude difference between target- and distractor-evoked activity predicts discrimination performance (rrm = 0.68). Specific adaptation of target or distractor shifts performance either below or above chance, respectively. It appears that recent auditory history profoundly influences stimulus saliency, biasing animals toward diametrically-opposed decisions.

Human listeners are able to recognize accurately an impressive range of complex sounds, such as musical instruments or voices. The underlying mechanisms are still poorly understood. Here, we aimed to characterize the processing time needed to recognize a natural sound. To do so, by analogy with the “rapid visual sequential presentation paradigm”, we embedded short target sounds within rapid sequences of distractor sounds. The core hypothesis is that any correct report of the target implies that sufficient processing for recognition had been completed before the time of occurrence of the subsequent distractor sound. We conducted four behavioral experiments using short natural sounds (voices and instruments) as targets or distractors. We report the effects on performance, as measured by the fastest presentation rate for recognition, of sound duration, number of sounds in a sequence, the relative pitch between target and distractors and target position in the sequence. Results showed a very rapid auditory recognition of natural sounds in all cases. Targets could be recognized at rates up to 30 sounds per second. In addition, the best performance was observed for voices in sequences of instruments. These results give new insights about the remarkable efficiency of timbre processing in humans, using an original behavioral paradigm to provide strong constraints on future neural models of sound recognition.

The auditory brainstem compares sound-evoked excitation and inhibition from both ears to compute sound source location. Although alterations to the anatomy and physiology of the auditory brainstem have been demonstrated in Fragile X Syndrome (FXS) it is not known whether these changes cause sound localization impairments in FXS. To test the hypothesis that FXS-related alterations to brainstem circuits impair spatial hearing abilities, a reflexive prepulse inhibition (PPI) task, with sound source location as the prepulse stimulus, was used to show that Fmr1 knockout mice have decreased inhibition of their startle responses. Specifically, Fmr1 mice show decreased PPI compared to wildtype during gap detection, changes in sound source location, and spatial release from masking with no alteration to their overall startle thresholds compared to wildtype. Lastly, Fmr1 mice have increased latency to respond in these tasks suggesting additional impairments in the pathway responsible for reacting to a startling sound.

Purpose Previous studies with children and adults have demonstrated a familiar talker advantage —better word recognition for familiar talkers. The goal of the current study was to test whether this phenomenon is modulated by a child's language ability. Method Sixty children with a range of language ability were trained to learn the voices of 3 foreign-accented, German–English bilingual talkers and received feedback about their performance. Both before and after this talker voice training, children completed a spoken word recognition task in which they heard consonant–vowel–consonant words mixed with noise that were spoken by the 3 familiarized talkers and by 3 unfamiliar German–English bilinguals. Results Two findings emerged from this study: First, children with both higher and lower language ability performed similarly on the familiarized talkers. Second, children with higher language scores performed similarly on both the familiarized and unfamiliar talkers, whereas children with lower language scores performed worse on the unfamiliar talkers compared to familiar talkers, suggesting an inability to generalize to novel, unfamiliar talkers who spoke with a similar accent. Discussion Together, these findings indicate that children with higher language scores are able to generalize knowledge about foreign-accented talkers to help spoken word recognition for novel talkers with the same accent. In contrast, children with lower language skills did not exhibit the same magnitude of generalization. This lack of generalization to similar talkers may mean that children with lower language skills are at a disadvantage in spoken language tasks because they are unable to process speech as well when listening to unfamiliar talkers.

Objectives: This study aimed to evaluate the informational component of speech-on-speech masking. Speech perception in the presence of a competing talker involves not only informational masking (IM) but also a number of masking processes involving interaction of masker and target energy in the auditory periphery. Such peripherally generated masking can be eliminated by presenting the target and masker in opposite ears (dichotically). However, this also reduces IM by providing listeners with lateralization cues that support spatial release from masking (SRM). In tonal sequences, IM can be isolated by rapidly switching the lateralization of dichotic target and masker streams across the ears, presumably producing ambiguous spatial percepts that interfere with SRM. However, it is not clear whether this technique works with speech materials. Design: Speech reception thresholds (SRTs) were measured in 17 young normal-hearing adults for sentences produced by a female talker in the presence of a competing male talker under three different conditions: diotic (target and masker in both ears), dichotic, and dichotic but switching the target and masker streams across the ears. Because switching rate and signal coherence were expected to influence the amount of IM observed, these two factors varied across conditions. When switches occurred, they were either at word boundaries or periodically (every 116 msec) and either with or without a brief gap (84 msec) at every switch point. In addition, SRTs were measured in a quiet condition to rule out audibility as a limiting factor. Results: SRTs were poorer for the four switching dichotic conditions than for the nonswitching dichotic condition, but better than for the diotic condition. Periodic switches without gaps resulted in the worst SRTs compared to the other switch conditions, thus maximizing IM. Conclusions: These findings suggest that periodically switching the target and masker streams across the ears (without gaps) was the most efficient in disrupting SRM. Thus, this approach can be used in experiments that seek a relatively pure measure of IM, and could be readily extended to translational research.

Partial hearing loss can cause neurons in the auditory and audiovisual cortices to increase their responsiveness to visual stimuli; however, behavioral studies in hearing-impaired humans and rats have found that the perceptual ability to accurately judge the relative timing of auditory and visual stimuli is largely unaffected. To investigate the neurophysiological basis of how audiovisual temporal acuity may be preserved in the presence of hearing loss-induced crossmodal plasticity, we exposed adult rats to loud noise and two weeks later performed in vivo electrophysiological recordings in two neighboring regions within the lateral extrastriate visual (V2L) cortex—a multisensory zone known to be responsive to audiovisual stimuli (V2L-Mz) and a predominantly auditory zone (V2L-Az). To examine the cortical layer-specific effects at the level of postsynaptic potentials, a current source density (CSD) analysis was applied to the local field potential (LFP) data recorded in response to auditory and visual stimuli presented at various stimulus onset asynchronies (SOAs). As predicted, differential effects were observed in the neighboring cortical regions’ postnoise exposure. Most notably, an analysis of the strength of multisensory response interactions revealed that V2L-Mz lost its sensitivity to the relative timing of the auditory and visual stimuli, due to an increased responsiveness to visual stimulation that produced a prominent audiovisual response irrespective of the SOA. In contrast, not only did the V2L-Az in noise-exposed rats become more responsive to visual stimuli but neurons in this region also inherited the capacity to process audiovisual stimuli with the temporal precision and specificity that was previously restricted to the V2L-Mz. Thus, the present study provides the first demonstration that audiovisual temporal processing can be preserved following moderate hearing loss via compensatory plasticity in the higher-order sensory cortices that is ultimately characterized by a functional transition in the cortical region capable of temporal sensitivity.

Objective: The purpose of this study was to investigate perceptual and electrophysiological encoding of complex periodic signals as a function of age. Design: Two groups of adults completed three listening tasks: a behavioural task of detection of a mistuned harmonic component in a complex tone, an electrophysiological measure of speech-evoked auditory brainstem response (sABR), and a speech-in-noise measure. Between group comparisons were undertaken for each task as well as pairwise correlation analyses for all tasks. Study sample: One group of younger adults (n = 20) and one group of older adults (n = 20) participated. All listeners had relatively normal audiometric thresholds (≤20 dB HL) from 250-4000 Hz. Results: Younger adults had better results than the older adults on all three tasks: sensitivity for detecting a mistuned harmonic, spectral encoding for sABR, and release from masking for the speech-in-noise test. There were no significant correlations between measures when evaluating the older adults in isolation. Conclusions: The results are consistent with the body of literature that demonstrates reduced temporal processing abilities for older adults. The combined method approach undertaken in this investigation did not result in correlations between the perceptual and electrophysiological measures of temporal processing.

A noise-induced loss of inner hair cell (IHC) - auditory nerve synaptic connections has been suggested as a factor that can trigger the progression of maladaptive plastic changes leading to noise-induced tinnitus. The present study used a military relevant small arms fire (SAF)-like noise (50 biphasic impulses over 2.5 min at 152 dB SPL given unilaterally to the right ear) to induce loss (∼1/3) of IHC synaptic ribbons (associated with synapse loss) in rat cochleae with only minor (less than 10%) loss of outer hair cells. Approximately half of the noise-exposed rats showed poorer Gap Detection post-noise, a behavioral indication suggesting the presence of tinnitus. There was significantly greater loss of IHC ribbons in noise-exposed rats with reduced Gap Detection compared to noise-exposed rats retaining normal Gap Detection. We have previously shown systemic administration of piribedil, memantine, and/or ACEMg significantly reduced loss of IHC ribbons induced by a 3 h 4 kHz octave band 117 dB (SPL) noise. The present study examined if this treatment would also reduce ribbon loss from the SAF-like noise exposure and if this would prevent the reduced Gap Detection. As in the previous study, piribedil, memantine, and ACEMg treatment significantly reduced the noise-induced loss of ribbons, such that it was no longer significantly different from normal. However, it did not prevent development of the reduced Gap Detection indication of tinnitus in all treated noise-exposed rats, reducing the incidence but not reaching significance.

To evaluate the ability of a restricted range of auditory-nerve fibers to encode a large perceptual dynamic range, Viemeister [(1983). Science 221, 1206–1208] examined the detection of a change in the level of a high-frequency band of noise flanked by more intense fixed-level noise maskers. Here, stimuli and procedures similar to Viemeister's were used, but random manipulations of level and notch cutoff frequency were included to evaluate predictions of energy-based models. The results indicate that cues other than the change in level per se are available, and suggest the potential contribution of changes in pitch/timbre for this task.

Previous studies suggest that envelope-following responses (EFRs) reveal important differences in temporal coding fidelity amongst listeners who have normal hearing thresholds, consistent with these listeners differing in the degree to which they suffer from cochlear synaptopathy. Like conventional hearing loss, the severity of cochlear synaptopathy may vary along the cochlea. A number of earlier studies have suggested methods for estimating EFRs driven by specific frequency regions of the cochlea, which would allow synaptopathy to be estimated as a function of cochlear place. Here, we tested a method for measuring EFRs from multiple locations along the cochlea simultaneously, using narrowband stimuli. We compared responses to multiple simultaneous narrowband complex harmonic tones in three non-overlapping frequency bands, each having a unique fundamental frequency, to responses to the individual narrowband stimuli alone, and to responses when noise was added to different combinations of the frequency bands. Our results suggest that simultaneous presentation of multiple tone complexes with different fundamental frequencies leads to repeatable measures of temporal coding fidelity at the cochlear frequency regions corresponding to the narrowband carrier frequencies. Other results suggested that while off-frequency contributions to EFRs driven by narrowband signals (due to spread of excitation) can add destructively to the on frequency response, these interactions were small compared to EFR magnitude. Overall, our results point to the utility of using multi-band complex tone stimuli to estimate the profile of temporal coding fidelity, and thus the degree of synaptopathy, as a function of cochlear place. This article is part of a Special Issue entitled: Hearing Loss, Tinnitus, Hyperacusis, Central Gain.

The central gain model of hyperacusis proposes that loss of auditory input can result in maladaptive neuronal gain increases in the central auditory system, leading to the over-amplification of sound-evoked activity and excessive loudness perception. Despite the attractiveness of this model, and supporting evidence for it, a critical test of the central gain theory requires that changes in sound-evoked activity be explicitly linked to perceptual alterations of loudness. Here we combined an operant conditioning task that uses a subject's reaction time to auditory stimuli to produce reliable measures of loudness growth with chronic electrophysiological recordings from the auditory cortex and inferior colliculus of awake, behaviorally-phenotyped animals. In this manner, we could directly correlate daily assessments of loudness perception with neurophysiological measures of sound encoding within the same animal. We validated this novel psychophysical-electrophysiological paradigm with a salicylate-induced model of hearing loss and hyperacusis, as high doses of sodium salicylate reliably induce temporary hearing loss, neural hyperactivity, and auditory perceptual disruptions like tinnitus and hyperacusis. Salicylate induced parallel changes to loudness growth and evoked response-intensity functions consistent with temporary hearing loss and hyperacusis. Most importantly, we found that salicylate-mediated changes in loudness growth and sound-evoked activity were correlated within individual animals. These results provide strong support for the central gain model of hyperacusis and demonstrate the utility of using an experimental design that allows for within-subject comparison of behavioral and electrophysiological measures, thereby making inter-subject variability a strength rather than a limitation.

Purpose As pulse rate increases beyond a few hundred Hertz, younger normal-hearing (NH) participants' ability to encode temporal information in band-limited acoustic pulse trains decreases, demonstrating a rate limitation in processing rapid temporal information. Rate discrimination abilities, however, have yet to be investigated in older NH participants—a population that experiences age-related temporal processing deficits. It was hypothesized that age-related temporal processing deficits lead to decreased temporal rate discrimination abilities in older compared with younger NH participants, which could be observed in both perceptual and electrophysiological measurements. Method Fifteen younger and 15 older NH participants were presented acoustic pulse trains with a 4-kHz center frequency and 1-kHz bandwidth at 75 dB SPL monaurally. The pulse rate was 80, 200, or 400 Hz. Just noticeable differences were obtained using an adaptive procedure that instructed the participants to identify the pulse train with the highest pitch. Auditory steady-state responses (ASSRs) were recorded to the same pulse trains with 2 additional rates—20 and 40 Hz. The Digit Symbol Coding and Digit Symbol Search subtests of the Wechsler Adult Intelligence Scale ( Wechsler, 1997 ) were measured as correlates to domain-general cognitive processing speed. Results As rate increased from 80 to 400 Hz, performance on the perceptual rate discrimination task worsened in both groups. ASSR spectral energy also decreased, but only in the older group. Perceptual performance was equivalent between groups across rates. The older group had lower ASSR spectral energy (lower signal-to-noise ratios) at the 400-Hz rate than the younger group, but there were no group differences for the other rates. The overall strength of neural rate representation, along with speed of processing performance, predicted perceptual performance for the 400-Hz rate. Conclusion These results suggest that neural representation at early levels of the auditory system and processing speed are factors in perceptual auditory temporal processing performance, especially in older adults.

Technology that translates neural activity into speech would be transformative for people who are unable to communicate as a result of neurological impairments. Decoding speech from neural activity is challenging because speaking requires very precise and rapid multi-dimensional control of vocal tract articulators. Here we designed a neural decoder that explicitly leverages kinematic and sound representations encoded in human cortical activity to synthesize audible speech. Recurrent neural networks first decoded directly recorded cortical activity into representations of articulatory movement, and then transformed these representations into speech acoustics. In closed vocabulary tests, listeners could readily identify and transcribe speech synthesized from cortical activity. Intermediate articulatory dynamics enhanced performance even with limited data. Decoded articulatory representations were highly conserved across speakers, enabling a component of the decoder to be transferrable across participants. Furthermore, the decoder could synthesize speech when a participant silently mimed sentences. These findings advance the clinical viability of using speech neuroprosthetic technology to restore spoken communication.

Purpose Age-related sensorineural hearing loss can dramatically affect speech recognition performance due to reduced audibility and suprathreshold distortion of spectrotemporal information. Normal aging produces changes within the central auditory system that impose further distortions. The goal of this study was to characterize the effects of aging and hearing loss on perceptual representations of speech. Method We asked whether speech intelligibility is supported by different patterns of spectrotemporal modulations (STMs) in older listeners compared to young normal-hearing listeners. We recruited 3 groups of participants: 20 older hearing-impaired (OHI) listeners, 19 age-matched normal-hearing listeners, and 10 young normal-hearing (YNH) listeners. Listeners performed a speech recognition task in which randomly selected regions of the speech STM spectrum were revealed from trial to trial. The overall amount of STM information was varied using an up–down staircase to hold performance at 50% correct. Ordinal regression was used to estimate weights showing which regions of the STM spectrum were associated with good performance (a “classification image” or CImg). Results The results indicated that (a) large-scale CImg patterns did not differ between the 3 groups; (b) weights in a small region of the CImg decreased systematically as hearing loss increased; (c) CImgs were also nonsystematically distorted in OHI listeners, and the magnitude of this distortion predicted speech recognition performance even after accounting for audibility; and (d) YNH listeners performed better overall than the older groups. Conclusion We conclude that OHI/older normal-hearing listeners rely on the same speech STMs as YNH listeners but encode this information less efficiently. Supplemental Material https://doi.org/10.23641/asha.7859981

Salience is a broad and widely used concept in neuroscience whose neuronal correlates, however, remain elusive. In behavioral conditioning, salience is used to explain various effects, such as stimulus overshadowing, and refers to how fast and strongly a stimulus can be associated with a conditioned event. Here, we identify sounds of equal intensity and perceptual detectability, which due to their spectro-temporal content recruit different levels of population activity in mouse auditory cortex. When using these sounds as cues in a Go/NoGo discrimination task, the degree of cortical recruitment matches the salience parameter of a reinforcement learning model used to analyze learning speed. We test an essential prediction of this model by training mice to discriminate light-sculpted optogenetic activity patterns in auditory cortex, and verify that cortical recruitment causally determines association or overshadowing of the stimulus components. This demonstrates that cortical recruitment underlies major aspects of stimulus salience during reinforcement learning.

Adults with bilateral cochlear implants (BiCIs) receive benefits in localizing stationary sounds when listening with two implants compared with one; however, sound localization ability is significantly poorer when compared to normal hearing (NH) listeners. Little is known about localizing sound sources in motion, which occurs in typical everyday listening situations. The authors considered the possibility that sound motion may improve sound localization in BiCI users by providing multiple places of information. Alternatively, the ability to compare multiple spatial locations may be compromised in BiCI users due to degradation of binaural cues, and thus result in poorer performance relative to NH adults. In this study, the authors assessed listeners' abilities to distinguish between sounds that appear to be moving vs stationary, and track the angular range and direction of moving sounds. Stimuli were bandpass-filtered (150–6000 Hz) noise bursts of different durations, panned over an array of loudspeakers. Overall, the results showed that BiCI users were poorer than NH adults in (i) distinguishing between a moving vs stationary sound, (ii) correctly identifying the direction of movement, and (iii) tracking the range of movement. These findings suggest that conventional cochlear implant processors are not able to fully provide the cues necessary for perceiving auditory motion correctly.

Information from different sensory modalities can interact, shaping what we think we have seen, heard, or otherwise perceived. Such interactions can enhance the precision of perceptual decisions, relative to those based on information from a single sensory modality. Several computational processes could account for such improvements. Slight improvements could arise if decisions are based on multiple independent sensory estimates, as opposed to just one. Still greater improvements could arise if initially independent estimates are summed to form a single integrated code. This hypothetical process has often been described as optimal when it results in bimodal performance consistent with a summation of unimodal estimates weighted in proportion to the precision of each initially independent sensory code. Here we examine cross-modal cue combination for audio-visual temporal rate and spatial location cues. While suggestive of a cross-modal encoding advantage, the degree of facilitation falls short of that predicted by a precision weighted summation process. These data accord with other published observations, and suggest that precision weighted combination is not a general property of human cross-modal perception.

Pitch perception is critical for recognizing speech, music and animal vocalizations, but its neurobiological basis remains unsettled, in part because of divergent results across species. We investigated whether species-specific differences exist in the cues used to perceive pitch and whether these can be accounted for by differences in the auditory periphery. Ferrets accurately generalized pitch discriminations to untrained stimuli whenever temporal envelope cues were robust in the probe sounds, but not when resolved harmonics were the main available cue. By contrast, human listeners exhibited the opposite pattern of results on an analogous task, consistent with previous studies. Simulated cochlear responses in the two species suggest that differences in the relative salience of the two pitch cues can be attributed to differences in cochlear filter bandwidths. The results support the view that cross-species variation in pitch perception reflects the constraints of estimating a sound’s fundamental frequency given species-specific cochlear tuning.

Auditory-nerve fibers are lost steadily with age and as a possible consequence of noise-induced glutamate excitotoxicity. Auditory-nerve loss in the absence of other cochlear pathologies is thought to be undetectable with a pure-tone audiogram while degrading real-world speech perception (hidden hearing loss). Perceptual deficits remain unclear, however, due in part to the limited behavioral capacity of existing rodent models to discriminate complex sounds. The budgerigar is an avian vocal learner with human-like behavioral sensitivity to many simple and complex sounds and the capacity to mimic speech. Previous studies in this species show that intracochlear kainic-acid infusion reduces wave 1 of the auditory brainstem response by 40-70%, consistent with substantial excitotoxic auditory-nerve damage. The present study used operant-conditioning procedures in trained budgerigars to quantify kainic-acid effects on tone detection across frequency (0.25-8 kHz; the audiogram) and as a function of duration (20-160 ms; temporal integration). Tone thresholds in control animals were lowest from 1 to 4 kHz and decreased with increasing duration as in previous studies of the budgerigar. Behavioral results in kainic-acid-exposed animals were as sensitive as in controls, suggesting preservation of the audiogram and temporal integration despite auditory-nerve loss associated with up to 70% wave 1 reduction. Distortion-product otoacoustic emissions were also preserved in kainic-acid exposed animals, consistent with normal hair-cell function. These results highlight considerable perceptual resistance of tone-detection performance with selective auditory-nerve loss. Future behavioral studies in budgerigars with auditory-nerve damage can use complex speech-like stimuli to help clarify aspects of auditory perception impacted by this common cochlear pathology.