Monaural Spectral Cues Research Articles

Sound localization acuity of the common marmoset (Callithrix jacchus)

The common marmoset (Callithrix jacchus) is a small arboreal New World primate which has emerged as a promising model in auditory neuroscience. One potentially useful application of this model system is in the study of the neural mechanism underlying spatial hearing in primate species, as the marmosets need to localize sounds to orient their head to events of interest and identify their vocalizing conspecifics that are not visible. However, interpretation of neurophysiological data on sound localization requires an understanding of perceptual abilities, and the sound localization behavior of marmosets has not been well studied. The present experiment measured sound localization acuity using an operant conditioning procedure in which marmosets were trained to discriminate changes in sound location in the horizontal (azimuth) or vertical (elevation) dimension. Our results showed that the minimum audible angle (MAA) for horizontal and vertical discrimination was 13.17° and 12.53°, respectively, for 2 to 32 kHz Gaussian noise. Removing the monaural spectral cues tended to increase the horizontal localization acuity (11.31°). Marmosets have larger horizontal MAA (15.54°) in the rear than the front. Removing the high-frequency (> 26 kHz) region of the head-related transfer function (HRTF) affected vertical acuity mildly (15.76°), but removing the first notch (12–26 kHz) region of HRTF substantially reduced the vertical acuity (89.01°). In summary, our findings indicate that marmosets’ spatial acuity is on par with other species of similar head size and field of best vision, and they do not appear to use monaural spectral cues for horizontal discrimination but rely heavily on first notch region of HRTF for vertical discrimination.

High-Frequency Hearing Is Required to Compute a Topographic Map of Auditory Space in the Mouse Superior Colliculus.

A topographic map of auditory space is a feature found in the superior colliculus (SC) of many species, including CBA/CaJ mice. In this genetic background, high-frequency monaural spectral cues and interaural level differences (ILDs) are used to compute spatial receptive fields (RFs) that form a topographic map along the azimuth. Unfortunately, C57BL/6 mice, a strain widely used for transgenic manipulation, display age-related hearing loss (AHL) because of an inbred mutation in the Cadherin 23 gene (Cdh23) that affects hair cell mechanotransduction. To overcome this problem, researchers have used young C57BL/6 mice in their studies, as they have been shown to have normal hearing thresholds. However, important details of the auditory response characteristics of the SC such as spectral responses and spatial localization, have not been characterized in young C57BL/6 mice. Here, we show that two- to four-month C57BL/6 mice lack neurons with frontal auditory RFs and therefore lack a topographic representation of auditory space in the SC. Analysis of the spectrotemporal RFs (STRFs) of the SC auditory neurons shows that C57BL/6 mouse SC neurons lack the ability to detect the high-frequency (>40 kHz) spectral cues that are needed to compute frontal RFs. We also show that crossing C57BL/6 mice with CBA/CaJ mice or introducing one copy of the wild-type Cdh23 to C57BL/6 mice rescues the high-frequency hearing deficit and improves the topographic map of auditory space. Taken together, these results demonstrate the importance of high-frequency hearing in computing a topographic map of auditory space.

Deep neural network models of sound localization reveal how perception is adapted to real-world environments

Mammals localize sounds using information from their two ears. Localization in real-world conditions is challenging, as echoes provide erroneous information, and noises mask parts of target sounds. To better understand real-world localization we equipped a deep neural network with human ears and trained it to localize sounds in a virtual environment. The resulting model localized accurately in realistic conditions with noise and reverberation. In simulated experiments, the model exhibited many features of human spatial hearing: sensitivity to monaural spectral cues and interaural time and level differences, integration across frequency, biases for sound onsets, and limits on localization of concurrent sources. But when trained in unnatural environments without either reverberation, noise, or natural sounds, these performance characteristics deviated from those of humans. The results show how biological hearing is adapted to the challenges of real-world environments and illustrate how artificial neural networks can reveal the real-world constraints that shape perception.

THE EFFECT OF FULL AND PARTIAL MAPPING ON SOUND LOCALIZATION AND CONSONANT RECOGNITION ABILITIES IN CI USERS

Sound localization is an auditory skill that allows identification of an acoustic signal in space. It’s dependent on two mechanisms; a peripheral mechanism that includes signal comparison between right and left ears of a normal hearing person and a central processing mechanism in the brain stem pathways.Hearing loss impairs both peripheral and central mechanisms. However, people with unilateral hearing loss have some ability to localize sound, a finding that reinforces the importance of monaural spectral cues for this purpose. Monaural ability to localize sound might be especially important for patients with unilateral cochlear implants, who base their ‘acoustic picture’ on limited acoustic data.Speech is a complex acoustic signal that is rich in both spectral and temporal features. The listener makes use of whatever spectral or temporal cues are available to help decode the incoming speech signal. Multiple redundant acoustic cues can contribute to the perception of a single phonemic contrast. CI processing preserves the slowly varying temporal envelope information in the speech signal (i.e., cues related to amplitude changes over time) but provides only a coarse representation of the spectral envelope.

Modeling perceived externalization of a static, lateral sound image

Perceived externalization is a relevant feature to create an immersive acoustic environment with headphone reproduction. In the present study, listener-specific acoustic transfer characteristics for an azimuth angle of 90° were modified to investigate the role of monaural spectral cues, interaural level differences (ILDs), and temporal fluctuations of ILDs on perceived externalization in anechoic and reverberant environments. Listeners’ ratings suggested that each acoustic cue was important for perceived externalization. If only one correct acoustic cue remained in the ear signals, the sound image could not be perceived as fully externalized. Reverberation did reduce but not eliminate the influences of monaural spectral and ILD cues on perceived externalization. Additionally, the spectral details of the ipsilateral ear signal were more important for perceived externalization than those in the contralateral ear signal. A computational model was proposed to quantify those relationships and predict externalization ratings by comparing the acoustic cues extracted from the target (modified) and template (non-processed) binaural signals after several auditory processing steps. The accuracy of predicted externalization ratings was higher than 90% under all experimental conditions.

Long-term evaluation of sound localisation in single-sided deaf adults fitted with a BAHA device.

To perform a long-term evaluation of the localisation capabilities in the horizontal plane of single-sided deaf patients fitted with a BAHA device. Single-centre retrospective study. Twenty-one adults with single-sided deafness (SSD) with normal hearing in the contralateral ear (pure tone average <20dB, SDS>90%) rehabilitated with a Cochlear BAHA device from 2003 to 2012 on the deaf side over a median follow-up of 8years. The task used in this paper is a sound localisation identification task with a set-up of seven loudspeakers on a semi-circular array at 30-degree intervals performed at three periods: before BAHA, initially and at last follow-up. Our main criterion of judgement was the root-mean-square (RMS) localisation error. In addition, the Bern Benefit in Single-Sided Deafness Questionnaire (BBSS) was administered. The mean RMS localisation error was initially estimated at 64° without any rehabilitation (for a chance level RMS estimated at 81°). Initially, with the BAHA device, the RMS localisation error dropped to 51°. At the last follow-up evaluation, a significant decrease at 23° was noted. Concerning the Bern Questionnaire, 19% of the patients (n=4) did not report any change (score of 0), 33% (n=7) are satisfied (score of +1 or +2) and 48% (n=10) are very satisfied with the BAHA device (score better than +3). Improvement of sound localisation in the horizontal plane for some SSD patients is likely related to altered processing of monaural spectral cues. The time needed to learn to use the azimuth-dependent spectral cues takes time. Long-term follow-up should be considered for studies investigating sound localisation performance.

The monaural spectral cues identified by a reverse correlation analysis of free-field auditory localization data.

The outer-ear's location-dependent pattern of spectral filtering generates cues used to determine a sound source's elevation as well as front-back location. The authors aim to identify these features using a reverse correlation analysis (RCA), combining free-field localization behaviour with the associated head-related transfer functions' (HRTFs) magnitude spectrum from a sample of 73 participants. Localization responses were collected before and immediately after introducing a pair of outer-ear inserts which modified the listener's HRTFs to varying extent. The RCA identified several different features responsible for eliciting localization responses. The efficacy of these was examined using two models of monaural localization. In general, the predicted performance was closely aligned with the free-field localization error for the bare-ear condition; however, both models tended to grossly over-estimate the localization error based on HRTFs modified by the outer-ear inserts. The RCA's feature selection notably had the effect of better aligning the predicted performance of both models to the actual localization performance. This suggests that the RCA revealed sufficient detail for both models to correctly predict localization performance and also limited the influence of filtered-out elements in the distorted HRTFs that contributed to the degraded accuracy of both models.

The Generalization of Auditory Accommodation to Altered Spectral Cues

The capacity of healthy adult listeners to accommodate to altered spectral cues to the source locations of broadband sounds has now been well documented. In recent years we have demonstrated that the degree and speed of accommodation are improved by using an integrated sensory-motor training protocol under anechoic conditions. Here we demonstrate that the learning which underpins the localization performance gains during the accommodation process using anechoic broadband training stimuli generalize to environmentally relevant scenarios. As previously, alterations to monaural spectral cues were produced by fitting participants with custom-made outer ear molds, worn during waking hours. Following acute degradations in localization performance, participants then underwent daily sensory-motor training to improve localization accuracy using broadband noise stimuli over ten days. Participants not only demonstrated post-training improvements in localization accuracy for broadband noises presented in the same set of positions used during training, but also for stimuli presented in untrained locations, for monosyllabic speech sounds, and for stimuli presented in reverberant conditions. These findings shed further light on the neuroplastic capacity of healthy listeners, and represent the next step in the development of training programs for users of assistive listening devices which degrade localization acuity by distorting or bypassing monaural cues.

Contralateral routing of signals disrupts monaural level and spectral cues to sound localisation on the horizontal plane

ObjectivesContra-lateral routing of signals (CROS) devices re-route sound between the deaf and hearing ears of unilaterally-deaf individuals. This rerouting would be expected to disrupt access to monaural level cues that can support monaural localisation in the horizontal plane. However, such a detrimental effect has not been confirmed by clinical studies of CROS use. The present study aimed to exercise strict experimental control over the availability of monaural cues to localisation in the horizontal plane and the fitting of the CROS device to assess whether signal routing can impair the ability to locate sources of sound and, if so, whether CROS selectively disrupts monaural level or spectral cues to horizontal location, or both. DesignUnilateral deafness and CROS device use were simulated in twelve normal hearing participants. Monaural recordings of broadband white noise presented from three spatial locations (−60°, 0°, and +60°) were made in the ear canal of a model listener using a probe microphone with and without a CROS device. The recordings were presented to participants via an insert earphone placed in their right ear. The recordings were processed to disrupt either monaural level or spectral cues to horizontal sound location by roving presentation level or the energy across adjacent frequency bands, respectively. Localisation ability was assessed using a three-alternative forced-choice spatial discrimination task. ResultsParticipants localised above chance levels in all conditions. Spatial discrimination accuracy was poorer when participants only had access to monaural spectral cues compared to when monaural level cues were available. CROS use impaired localisation significantly regardless of whether level or spectral cues were available. For both cues, signal re-routing had a detrimental effect on the ability to localise sounds originating from the side of the deaf ear (−60°). CROS use also impaired the ability to use level cues to localise sounds originating from straight ahead (0°). ConclusionsThe re-routing of sounds can restrict access to the monaural cues that provide a basis for determining sound location in the horizontal plane. Perhaps encouragingly, the results suggest that both monaural level and spectral cues may not be disrupted entirely by signal re-routing and that it may still be possible to reliably identify sounds originating on the hearing side.

Modeling the Effects of Sensorineural Hearing Loss on Sound Localization in the Median Plane.

Listeners use monaural spectral cues to localize sound sources in sagittal planes (along the up-down and front-back directions). How sensorineural hearing loss affects the salience of monaural spectral cues is unclear. To simulate the effects of outer-hair-cell (OHC) dysfunction and the contribution of different auditory-nerve fiber types on localization performance, we incorporated a nonlinear model of the auditory periphery into a model of sagittal-plane sound localization for normal-hearing listeners. The localization model was first evaluated in its ability to predict the effects of spectral cue modifications for normal-hearing listeners. Then, we used it to simulate various degrees of OHC dysfunction applied to different types of auditory-nerve fibers. Predicted localization performance was hardly affected by mild OHC dysfunction but was strongly degraded in conditions involving severe and complete OHC dysfunction. These predictions resemble the usually observed degradation in localization performance induced by sensorineural hearing loss. Predicted localization performance was best when preserving fibers with medium spontaneous rates, which is particularly important in view of noise-induced hearing loss associated with degeneration of this fiber type. On average across listeners, predicted localization performance was strongly related to level discrimination sensitivity of auditory-nerve fibers, indicating an essential role of this coding property for localization accuracy in sagittal planes.

Modeling Localization of Amplitude-Panned Virtual Sources in Sagittal Planes.

Vector-base amplitude panning (VBAP) aims at creating virtual sound sources at arbitrary directions within multichannel sound reproduction systems. However, VBAP does not consistently produce listener-specific monaural spectral cues that are essential for localization of sound sources in sagittal planes, including the front-back and up-down dimensions. In order to better understand the limitations of VBAP, a functional model approximating human processing of spectro-spatial information was applied to assess accuracy in sagittal-plane localization of virtual sources created by means of VBAP. First, we evaluated VBAP applied on two loudspeakers in the median plane, and then we investigated the directional dependence of the localization accuracy in several three-dimensional loudspeaker arrangements designed in layers of constant elevation. The model predicted a strong dependence on listeners' individual head-related transfer functions, on virtual source directions, and on loudspeaker arrangements. In general, the simulations showed a systematic degradation with increasing polar-angle span between neighboring loudspeakers. For the design of VBAP systems, predictions suggest that spans up to 40° polar angle yield a good trade-off between system complexity and localization accuracy. Special attention should be paid to the frontal region where listeners are most sensitive to deviating spectral cues.

Combination of two consecutive monaural “looks” as a spatial hearing cue

Interaural time, level, and spectral differences are the major cues being used for sound source localization in the horizontal plane. Past studies have shown that human subjects could be trained to use monaural spectral cues to localize the sources of sound in the azimuth plane, but performance is poor. The current study investigates whether the combination of two monaural signals at both ears, one after another in time, could benefit sound source localization accuracy. The purpose is to study the possibility that the human subjects can compare a monaural signal at one ear to the short term memory of a prior monaural signal arriving at the other ear. Subjects were asked to judge the position of a loudspeaker that presented a 250-ms, 40 dBA noise burst with a roving spectral contour in a quarter field. The rms sound source localization error was measured in three conditions: (1) a single monaural signal; (2) two consecutive monaural “looks” at identical signals, with an interval of 3 s and (3) normal binaural hearing. It was found that the “two looks” localization performance was better than that in the case of monaural presentation, but is still inferior to that of the binaural presentation. Similar experiments were carried out over headphones. Lateralization performance was compared to the sound-field localization data. Contributions of level and spectral differences will be discussed. [Research supported by the AFOSR.]

Accommodating to new ears: the effects of sensory and sensory-motor feedback.

Changing the shape of the outer ear using small in-ear molds degrades sound localization performance consistent with the distortion of monaural spectral cues to location. It has been shown recently that adult listeners re-calibrate to these new spectral cues for locations both inside and outside the visual field. This raises the question as to the teacher signal for this remarkable functional plasticity. Furthermore, large individual differences in the extent and rate of accommodation suggests a number of factors may be driving this process. A training paradigm exploiting multi-modal and sensory-motor feedback during accommodation was examined to determine whether it might accelerate this process. So as to standardize the modification of the spectral cues, molds filling 40% of the volume of each outer ear were custom made for each subject. Daily training sessions for about an hour, involving repetitive auditory stimuli and exploratory behavior by the subject, significantly improved the extent of accommodation measured by both front-back confusions and polar angle localization errors, with some improvement in the rate of accommodation demonstrated by front-back confusion errors. This work has implications for both the process by which a coherent representation of auditory space is maintained and for accommodative training for hearing aid wearers.

Relearning Auditory Spectral Cues for Locations Inside and Outside the Visual Field

Previous research has demonstrated that, over a period of weeks, the auditory system accommodates to changes in the monaural spectral cues for sound locations within the frontal region of space. We were interested to determine if similar accommodation could occur for locations in the posterior regions of space, i.e. in the absence of contemporaneous visual information that indicates any mismatch between the perceived and actual location of a sound source. To distort the normal spectral cues to sound location, eight listeners wore small moulds in each ear. HRTF recordings confirmed that while the moulds substantially altered the monaural spectral cues, sufficient residual cues were retained to provide a basis for relearning. Compared to control measures, sound localization performance initially decreased significantly, with a sevenfold increase in front-back confusions and elevation errors more than doubled. Subjects wore the moulds continuously for a period of up to 60 days (median 38 days), over which time performance improved but remained significantly poorer than control levels. Sound localization performance for frontal locations (audio-visual field) was compared with that for posterior space (audio-only field), and there was no significant difference between regions in either the extent or rate of accommodation. This suggests a common mechanism for both regions of space that does not rely on contemporaneous visual information as a teacher signal for recalibration of the auditory system to modified spectral cues.

Aging Effects on Detection of Spectral Changes Induced by a Break in Sound Correlation

Previous studies have shown that both younger adults and older adults with clinically normal hearing are able to detect a break in correlation (BIC) between interaurally correlated sounds presented over headphones. This ability to detect a BIC improved when the correlated sounds were presented over left and right loudspeakers rather than over left and right headphones, suggesting that additional spectral cues provided by comb filtering (caused by interference between the two channels) facilitate detection of the BIC. However, older adults receive significantly less benefit than younger adults from a switch to loudspeaker presentation. It is hypothesized that this is a result of an age-related reduction in the sensitivity to the monaural spectral cues provided by comb filtering. Two experiments were conducted in this study. Correlated white noises with a BIC in the temporal middle were presented from two spatially separated loudspeakers (positioned at ±45-degree azimuth) and recorded at the right ear of a Knowles Electronic Manikin for Acoustic Research (KEMAR). In Experiment 1, the waveforms recorded at the KEMAR's right ear were presented to the participant's right ear over a headphone in 14 younger adults and 24 older adults with clinically normal hearing. In Experiment 2, 8 of the 14 younger participants participated. Under the monaurally cueing condition, the waveforms recorded at the KEMAR's right ear were presented to the participant's right ear as Experiment 1. Under the binaurally cueing condition, waveforms delivered from the left loudspeaker and those from the right loudspeaker were recorded at the KEMAR's left and right ear, respectively, thereby eliminating the spectral ripple cue, and were presented to the participant's left and right ears, respectively. For each of the two experiments, the break duration threshold for detecting the BIC was examined when the interloudspeaker interval (delay) (ILI) was 0, 1, 2, or 4 msec (left loudspeaker leading). In Experiment 1, both younger participants and older participants detected the BIC in the waveforms recorded by the right ear of KEMAR, but older participants had higher detection thresholds than younger participants when the ILI was 0, 2, or 4 msec without an effect of SPL shift between 59 and 71 dB. In Experiment 2, each of the eight younger participants was able to detect the occurrence of the BIC in either the monaurally cueing or binaural-cueing condition. In addition, the detection threshold under the monaurally cueing condition was substantially the same as that under the binaurally cueing condition at each of the four ILIs. Younger adults and older adults with clinically normal hearing are able to detect the monaural spectral changes arising from comb filtering when a sudden drop in intersound correlation is introduced. However, younger adults are more sensitive than older adults are, at detecting the BIC. The findings suggest that older adults are less able than younger adults to detect a periodic ripple in the sound spectrum. This age-related ability reduction may contribute to older adults' difficulties in hearing under noisy, reverberant conditions.

Detection of spectral changes induced by a break in sound correlation in younger adults and older adults

Detecting a transient break in correlation (BIC) between correlated sounds is much easier when presented over two loudspeakers than when presented over two headphones. However, older adults benefit less than younger adults from a change from headphone to loudspeaker presentation (Ear and Hearing, (30) 273-286, 2009), suggesting an age-related reduction in sensitivity to monaural and/or binaural spectral cues provided by comb filtering. In this study, the monaural spectral cues present in the sound field were isolated and extracted, and then presented over headphones to younger adults and older adults with clinically normal hearing. Compared to younger adults, older adults exhibited a reduced sensitivity to the monaural spectral cues, particularly when an inter-loudspeaker time interval was introduced.

Soil erosion hazard maps for corn stover management using National Resources Inventory data and the Water Erosion Prediction Project

A topographic map of auditory space is a feature found in the superior colliculus (SC) of many species, including CBA/CaJ mice. In this genetic background, high-frequency monaural spectral cues and interaural level differences are used to compute spatial receptive fields (RFs) that form a topographic map along the azimuth. Unfortunately, C57BL/6 mice, a strain widely used for transgenic manipulation, display age-related hearing loss (AHL) due to an inbred mutation in the Cadherin 23 gene (<i>Cdh23)</i> that affects hair cell mechanotransduction. To overcome this problem, researchers have used young C57BL/6 mice in their studies, as they have been shown to have normal hearing thresholds. However, important details of the auditory response characteristics of the SC such as spectral responses and spatial localization, have not been characterized in young C57BL/6 mice. Here we show that 2-4-month C57BL/6 mice lack neurons with frontal auditory RFs and therefore lack a topographic representation of auditory space in the SC. Analysis of the spectrotemporal receptive fields (STRFs) of the SC auditory neurons shows that C57BL/6 mouse SC neurons lack the ability to detect the high-frequency (&gt;40kHz) spectral cues that are needed to compute frontal RFs. We also show that crossing C57BL/6 mice with CBA/CaJ mice or introducing one copy of the wild-type <i>Cdh23</i> to C57BL/6 mice rescues the high-frequency hearing deficit and improves the topographic map of auditory space. Taken together, these results demonstrate the importance of high-frequency hearing in computing a topographic map of auditory space. <b>Significance Statement</b> Despite the strain’s age-dependent hearing loss, C57BL/6 mice are widely used in auditory studies because of the development of transgenic reporter and Cre lines in this genetic background. Here we examined the topographic map of auditory space and spectrotemporal properties of neurons in the SC of C57BL/6 mice. We found an early-onset high-frequency hearing deficit that results in the loss of SC neurons with frontal RFs and, consequently, an absence of a topographic map of auditory space. These findings stress the importance of high-frequency hearing to compute spatially restricted receptive fields and serve as a caution to researchers that doing auditory-related research using the C57BL/6 genetic background may not be representative of true wild-type mice.

Adaptive Reweighting of Auditory Localization Cues in Response to Chronic Unilateral Earplugging in Humans

Localizing a sound source involves the detection and integration of various spatial cues present in the sound waves at each ear. Previous studies indicate that the brain circuits underlying sound localization are calibrated by experience of the cues available to each individual. Plasticity in spatial hearing is most pronounced during development but can also be demonstrated during adulthood under certain circumstances. Investigations into whether adult humans can adjust to reduced input in one ear and learn a new correspondence between interaural differences cues and directions in space have produced conflicting results. Here we show that humans of both sexes can relearn to localize broadband sounds with a flat spectrum in the horizontal plane after altering the spatial cues available by plugging one ear. In subjects who received daily training, localization accuracy progressively shifted back toward their pre-plug performance after 1 week of earplugging, whereas no improvement was seen if all trials were performed on the same day. However, localization performance did not improve on a task that used stimuli in which the source spectrum was randomized from trial to trial, indicating that monaural spectral cues are needed for plasticity. We also characterized the effects of the earplug on sensitivity to interaural time and level differences and found no clear evidence for adaptation to these cues as the free-field localization performance improved. These findings suggest that the mature auditory system can accommodate abnormal inputs and maintain a stable spatial percept by reweighting different cues according to how informative they are.

Shared Cortical Systems for Processing of Horizontal and Vertical Sound Motion

Cortical processing of horizontal and vertical sound motion in free-field space was investigated using high-density electroencephalography in combination with standardized low-resolution brain electromagnetic tomography (sLORETA). Eighteen subjects heard sound stimuli that, after an initial stationary phase in a central position, started to move centrifugally, either to the left, to the right, upward, or downward. The delayed onset of both horizontal and vertical motion elicited a specific motion-onset response (MOR), resulting in widely distributed activations, with prominent maxima in primary and nonprimary auditory cortices, insula, and parietal lobe. The comparison of MORs to horizontal and vertical motion orientations did not indicate any significant differences in latency or topography. Contrasting the sLORETA solutions for the two motion orientations revealed only marginal activation in postcentral gyrus. These data are consistent with the notion that azimuth and elevation components of dynamic auditory spatial information are processed in common, rather than separate, cortical substrates. Furthermore, the findings support the assumption that the MOR originates at a stage of auditory analysis after the different spatial cues (interaural and monaural spectral cues) have been integrated into a unified space code.

Event-related potentials for interaural time differences and spectral cues

Dichotic pitches and mistuned harmonics can each lead to the perception of one or two auditory objects. Comparison of event-related potentials for the perception of one versus two objects reveals an early negative and a late positive component. The relationship of these components with auditory segregation was further investigated using stimuli containing monaural spectral cues to pitch, binaural timing cues to pitch, or a combination of both, interleaved with control stimuli (no pitch). Stimuli containing timing cues or a combination of timing and spectral cues reliably elicited both components, which were of larger amplitude when both cues were present. For stimuli containing only spectral cues, the early component was attenuated in amplitude and no measurable late component was detected.