Representation Of Pitch Research Articles

The effect of stimulus context on pitch representations in the human auditory cortex

Neuroimaging studies of pitch coding seek to identify pitch-related responses separate from responses to other properties of the stimulus, such as its energy onset, and other general aspects of the listening context. The current study reports the first attempt to evaluate these modulatory influences using functional magnetic resonance imaging (fMRI) measures of cortical pitch representations. Stimulus context was manipulated using a ‘classical stimulation paradigm’ (whereby successive pitch stimuli were separated by gaps of silence) and a ‘continuous stimulation paradigm’ (whereby successive pitch stimuli were interspersed with noise to maintain a stable envelope). Pitch responses were measured for two types of pitch-evoking stimuli; a harmonic-complex tone and a complex Huggins pitch. Results for a group of 15 normally hearing listeners revealed that context effects were mostly observed in primary auditory regions, while the most significant pitch responses were localized to posterior nonprimary auditory cortex, specifically planum temporale. Sensitivity to pitch was greater for the continuous stimulation conditions perhaps because they better controlled for concurrent responses to the noise energy onset and reduced the potential problem of a non-linear fMRI response becoming saturated. These results provide support for hierarchical processing within human auditory cortex, with some parts of primary auditory cortex engaged by general auditory energy, some parts of planum temporale specifically responsible for representing pitch information and adjacent regions that are responsible for complex higher-level auditory processing such as representing pitch information as a function of listening context.

A novel physiological mechanism for pitch encoding in monkey primary auditory cortex.

Pitch is a fundamental perceptual attribute of hearing. While auditory cortex is implicated in pitch perception, how pitch is represented at the cortical level remains unclear. The present study examines a novel hypothesis for how the pitch of pure tones and of harmonic complex tones, with or without the fundamental frequency, is encoded in primary auditory cortex (A1): pitch is represented non‐topographically by the temporal distribution of population activity in A1. Sounds of lower pitch evoke a greater proportion of sustained multiunit activity (MUA), relative to initial onset MUA, than sounds of higher pitch, such that the temporal distribution of MUA systematically varies with pitch. Pure tones and harmonic complexes with the same pitch evoke a similar proportion of sustained MUA. The temporal distribution of MUA is largely invariant to changes in stimulus parameters (e.g., level and relative phase) that leave the perceived pitch unchanged. Timing of AEP components recorded in superficial layers of A1 parallels similar pitch‐related changes in AEPs recorded in humans. Coding of perceptual qualities based on the time course of neural activity has been proposed in other sensory modalities (e.g., olfaction) and offers a novel alternative to topographic representations of pitch at the cortical level.

Neural Coding of Periodicity in Marmoset Auditory Cortex

Pitch, our perception of how high or low a sound is on a musical scale, crucially depends on a sound's periodicity. If an acoustic signal is temporally jittered so that it becomes aperiodic, the pitch will no longer be perceivable even though other acoustical features that normally covary with pitch are unchanged. Previous electrophysiological studies investigating pitch have typically used only periodic acoustic stimuli, and as such these studies cannot distinguish between a neural representation of pitch and an acoustical feature that only correlates with pitch. In this report, we examine in the auditory cortex of awake marmoset monkeys (Callithrix jacchus) the neural coding of a periodicity's repetition rate, an acoustic feature that covaries with pitch. We first examine if individual neurons show similar repetition rate tuning for different periodic acoustic signals. We next measure how sensitive these neural representations are to the temporal regularity of the acoustic signal. We find that neurons throughout auditory cortex covary their firing rate with the repetition rate of an acoustic signal. However, similar repetition rate tuning across acoustic stimuli and sensitivity to temporal regularity were generally only observed in a small group of neurons found near the anterolateral border of primary auditory cortex, the location of a previously identified putative pitch processing center. These results suggest that although the encoding of repetition rate is a general component of auditory cortical processing, the neural correlate of periodicity is confined to a special class of pitch-selective neurons within the putative pitch processing center of auditory cortex.

The representation of pitch in auditory imagery: Evidence from S-R compatibility and distance effects

The representation of pitch in auditory imagery: Evidence from S-R compatibility and distance effects

The representation of pitch in auditory imagery: Evidence from S-R compatibility and distance effects

In three experiments we explored the nature of representations constructed during the perception and imagination of pitch. We employed a same–different task to eliminate the influence of nonauditory information and to minimise use of cognitive strategies on auditory imagery. A reference tone of frequency 1000, 1500, or 2000 Hz, or an imagined tone of a pitch indicated by a visual cue, was followed by a comparison tone (1000, 1500, or 2000 Hz) to which either a speeded same or different response was required. In separate experiments, same–different judgements were mapped to vertically (Experiments 1 and 2) and horizontally arranged responses (Experiment 3). Judgements of tones closer in pitch yielded longer reaction times and higher error rates than more distant tones, indicating a pitch distance effect for perceptual and imagery tasks alike. In addition, in the imagery task, same–different responses were faster when low-pitched tones demanded a bottom or left key response and high-pitched tones a top or right response than vice versa, suggesting that pitch is coded spatially. Together, these behavioural effects support the assumption that both perceived and imagined pitch are translated into an analogical representation in the spatial domain.

Free will debates: Simple experiments are not so simple.

Congenital amusia (amusia, hereafter) is a developmental disorder that impacts negatively on the perception of music. Psychophysical testing suggests that individuals with amusia have above average thresholds for detection of pitch change and pitch direction discrimination; however, a low-level auditory perceptual problem cannot completely explain the disorder, since discrimination of melodies is also impaired when the constituent intervals are suprathreshold for perception. The aim of the present study was to test pitch memory as a function of (a) time and (b) tonal interference, in order to determine whether pitch traces are inherently weaker in amusic individuals. Memory for the pitch of single tones was compared using two versions of a paradigm developed by Deutsch (1970a). In both tasks, participants compared the pitch of a standard (S) versus a comparison (C) tone. In the time task, the S and C tones were presented, separated in time by 0, 1, 5, 10, and 15 s (blocked presentation). In the interference task, the S and C tones were presented with a fixed time interval (5 s) but with a variable number of irrelevant tones in between 0, 2, 4, 6, and 8 tones (blocked presentation). In the time task, control performance remained high for all time intervals, but amusics showed a performance decrement over time. In the interference task, controls and amusics showed a similar performance decrement with increasing number of irrelevant tones. Overall, the results suggest that the pitch representations of amusic individuals are less stable and more prone to decay than those of matched non-amusic individuals.

Brainstem pitch representation in native speakers of Mandarin is less susceptible to degradation of stimulus temporal regularity

It has been demonstrated that neural encoding of pitch in the auditory brainstem is shaped by long-term experience with language. To date, however, all stimuli have exhibited a high degree of pitch saliency. The experimental design herein permits us to determine whether experience-dependent pitch representation in the brainstem is less susceptible to progressive degradation of the temporal regularity of iterated rippled noise (IRN). Brainstem responses were recorded from Chinese and English participants in response to IRN homologues of Mandarin Tone 2 (T2 IRN). Six different iterations steps were utilized to systematically vary the degree of temporal regularity in the fine structure of the IRN stimuli to produce a pitch salience continuum ranging from low to high. Pitch-tracking accuracy and pitch strength were computed from the brainstem responses using autocorrelation algorithms. Analysis of variance of brainstem responses to T2 IRN revealed that pitch-tracking accuracy is higher in the native tone language group (Chinese) relative to the non-tone language group (English) except for the three lowest steps along the continuum, and moreover, that pitch strength is greater in the Chinese group even in severely degraded stimuli for two of the six 40-ms sections of T2 IRN that exhibit rapid changes in pitch. For these same two sections, exponential time constants for the stimulus continuum revealed that pitch strength emerges 2–3 times faster in the tone language than in the non-tone language group as a function of increasing pitch salience. These findings altogether suggest that experience-dependent brainstem mechanisms for pitch are especially sensitive to those dimensions of tonal contours that provide cues of high perceptual saliency in degraded as well as normal listening conditions.

The effects of tone language experience on pitch processing in the brainstem

Neural encoding of pitch in the auditory brainstem is shaped by long-term experience with language. The aim herein was to determine to what extent this experience-dependent effect is specific to a particular language. Analysis of variance of brainstem responses to Mandarin and Thai tones revealed that regardless of language identity, pitch-tracking accuracy of whole tones was higher in the two tone language groups (Chinese, Thai) compared to the non-tone language group (English), and that pitch strength of 40-ms tonal sections was generally more robust in tone relative to non-tone languages. Discriminant analysis of tonal sections, as defined by variation in direction and degree of slope, showed that moderate rising pitch was the most important variable for classifying English, Chinese, and Thai participants into their respective groups. We conclude that language-dependent enhancement of pitch representation transfers to other languages with similar phonological systems. From a neurobiological perspective, these findings suggest that neural mechanisms local to the brainstem are tuned for processing pitch dimensions that are perceptually salient depending upon the melodic patterns of a language.

Dichotic pitch activates pitch processing centre in Heschl's gyrus

Although several neuroimaging studies have reported pitch-evoked activations at the lateral end of Heschl's gyrus, it is still under debate whether these findings truly represent activity in relation to the perception of pitch or merely stimulus-related features of pitch-evoking sounds. We investigated this issue in a functional magnetic resonance imaging (fMRI) experiment using pure tones in noise and dichotic pitch sequences, which either contained a melody or a fixed pitch. Dichotic pitch evokes a sensation of pitch only in binaural listening conditions, while the monaural signal cannot be distinguished from random noise. Our data show similar neural activations for both tones in noise and dichotic pitch, which are perceptually similar, but physically different. Pitch-related activation was found at the lateral end of Heschl's gyrus in both hemispheres, providing new evidence for a general involvement of this region in pitch processing. In line with prior studies, we found melody-related activation in Planum temporale and Planum polare, but not in primary auditory areas. These results support the view of a general representation of pitch in auditory cortex, irrespective of the physical attributes of the pitch-evoking sound.

Experience‐induced Malleability in Neural Encoding of Pitch, Timbre, and Timing

Speech and music are highly complex signals that have many shared acoustic features. Pitch, Timbre, and Timing can be used as overarching perceptual categories for describing these shared properties. The acoustic cues contributing to these percepts also have distinct subcortical representations which can be selectively enhanced or degraded in different populations. Musically trained subjects are found to have enhanced subcortical representations of pitch, timbre, and timing. The effects of musical experience on subcortical auditory processing are pervasive and extend beyond music to the domains of language and emotion. The sensory malleability of the neural encoding of pitch, timbre, and timing can be affected by lifelong experience and short-term training. This conceptual framework and supporting data can be applied to consider sensory learning of speech and music through a hearing aid or cochlear implant.

Finding the Pitch of the Missing Fundamental in Infants

Pitch perception is critical for the perception of speech and music, for object identification, and for auditory scene analysis, whereby representations are derived for each sounding object in the environment from the complex sound wave that reaches the ears. The perceived pitch of a complex sound corresponds to its fundamental frequency. However, removal of energy at the fundamental does not alter the pitch because adults use the harmonics to derive the pitch (Bendor and Wang, 2005; Trainor, 2008). Although sound frequency is represented subcortically, the integration of harmonics into a representation of pitch does not occur until auditory cortex (Bendor and Wang, 2005). Given that auditory cortex is immature in young infants, we examined the development of cortical representations for pitch by measuring electrophysiological (EEG) responses to pitch changes that required processing the pitch of the missing fundamental. Adults and infants 4 months and older showed a mismatch negativity response to these pitch changes, but 3-month-old infants did not. Thus, cortical representations of the pitch of the missing fundamental emerge between 3 and 4 months of age, indicating that there is a profound change in auditory perception for pitch in early infancy.

Experience-dependent Enhancement of Linguistic Pitch Representation in the Brainstem Is Not Specific to a Speech Context

Neural representation of pitch is influenced by lifelong experiences with music and language at both cortical and subcortical levels of processing. The aim of this article is to determine whether neural plasticity for pitch representation at the level of the brainstem is dependent upon specific dimensions of pitch contours that commonly occur as part of a native listener's language experience. Brainstem frequency following responses (FFRs) were recorded from Chinese and English participants in response to four Mandarin tonal contours presented in a nonspeech context in the form of iterated rippled noise. Pitch strength (whole contour, 250 msec; 40-msec segments) and pitch-tracking accuracy (whole contour) were extracted from the FFRs using autocorrelation algorithms. Narrow band spectrograms were used to extract spectral information. Results showed that the Chinese group exhibits smoother pitch tracking than the English group in three out of the four tones. Moreover, cross-language comparisons of pitch strength of 40-msec segments revealed that the Chinese group exhibits more robust pitch representation of those segments containing rapidly changing pitch movements across all four tones. FFR spectral data were complementary showing that the Chinese group exhibits stronger representation of multiple pitch-relevant harmonics relative to the English group across all four tones. These findings support the view that at early preattentive stages of subcortical processing, neural mechanisms underlying pitch representation are shaped by particular dimensions of the auditory stream rather than speech per se. Adopting a temporal correlation analysis scheme for pitch encoding, we propose that long-term experience sharpens the tuning characteristics of neurons along the pitch axis with enhanced sensitivity to linguistically relevant variations in pitch.

Spatial Representations Are Specific to Different Domains of Knowledge

There is evidence that many abstract concepts are represented cognitively in a spatial format. However, it is unknown whether similar spatial processes are employed in different knowledge domains, or whether individuals exhibit similar spatial profiles within and across domains. This research investigated similarities in spatial representation in two knowledge domains – mathematics and music. Sixty-one adults completed analogous number magnitude and pitch discrimination tasks: the Spatial-Numerical Association of Response Codes and Spatial-Musical Association of Response Codes tasks. Subgroups of individuals with different response patterns were identified through cluster analyses. For both the mathematical and musical tasks, approximately half of the participants showed the expected spatial judgment effect when explicitly cued to focus on the spatial properties of the stimuli. Despite this, performances on the two tasks were largely independent. Consistent with previous research, the study provides evidence for the spatial representation of number and pitch in the majority of individuals. However, there was little evidence to support the claim that the same spatial representation processes underpin mathematical and musical judgments.

Learning to Encode Timing: Mechanisms of Plasticity in the Auditory Brainstem

Mechanisms of plasticity have traditionally been ascribed to higher-order sensory processing areas such as the cortex, whereas early sensory processing centers have been considered largely hard-wired. In agreement with this view, the auditory brainstem has been viewed as a nonplastic site, important for preserving temporal information and minimizing transmission delays. However, recent groundbreaking results from animal models and human studies have revealed remarkable evidence for cellular and behavioral mechanisms for learning and memory in the auditory brainstem.

The amusic brain: in tune, out of key, and unaware

Like language, music engagement is universal, complex and present early in life. However, approximately 4% of the general population experiences a lifelong deficit in music perception that cannot be explained by hearing loss, brain damage, intellectual deficiencies or lack of exposure. This musical disorder, commonly known as tone-deafness and now termed congenital amusia, affects mostly the melodic pitch dimension. Congenital amusia is hereditary and is associated with abnormal grey and white matter in the auditory cortex and the inferior frontal cortex. In order to relate these anatomical anomalies to the behavioural expression of the disorder, we measured the electrical brain activity of amusic subjects and matched controls while they monitored melodies for the presence of pitch anomalies. Contrary to current reports, we show that the amusic brain can track quarter-tone pitch differences, exhibiting an early right-lateralized negative brain response. This suggests near-normal neural processing of musical pitch incongruities in congenital amusia. It is important because it reveals that the amusic brain is equipped with the essential neural circuitry to perceive fine-grained pitch differences. What distinguishes the amusic from the normal brain is the limited awareness of this ability and the lack of responsiveness to the semitone changes that violate musical keys. These findings suggest that, in the amusic brain, the neural pitch representation cannot make contact with musical pitch knowledge along the auditory-frontal neural pathway.

Experience-dependent neural representation of dynamic pitch in the brainstem

Brainstem frequency-following responses were recorded from Chinese and English participants in response to an iterated rippled noise homologue of Mandarin Tone 2 (T2) and linear and inverted curvilinear variants. Pitch-tracking accuracy and pitch strength analyses showed advantages for the Chinese group over the English in response to T2 only. Pitch strength was larger for the Chinese group in rapidly changing sections of T2 compared with corresponding sections of a linear ramp. We conclude that experience-dependent neural plasticity at subcortical levels of representation is highly sensitive to specific features of pitch patterns in one's native language. Such experience-dependent effects suggest that subcortical sensory encoding interacts with cognitive processing in the cerebral cortex to shape the perceptual system's response to pitch patterns.

Interdependent encoding of pitch, timbre, and spatial location in auditory cortex.

Because we can perceive the pitch, timbre, and spatial location of a sound source independently, it seems natural to suppose that cortical processing of sounds might separate out spatial from nonspatial attributes. Indeed, recent studies support the existence of anatomically segregated "what" and "where" cortical processing streams. However, few attempts have been made to measure the responses of individual neurons in different cortical fields to sounds that vary simultaneously across spatial and nonspatial dimensions. We recorded responses to artificial vowels presented in virtual acoustic space to investigate the representations of pitch, timbre, and sound source azimuth in both core and belt areas of ferret auditory cortex. A variance decomposition technique was used to quantify the way in which altering each parameter changed neural responses. Most units were sensitive to two or more of these stimulus attributes. Although indicating that neural encoding of pitch, location, and timbre cues is distributed across auditory cortex, significant differences in average neuronal sensitivity were observed across cortical areas and depths, which could form the basis for the segregation of spatial and nonspatial cues at higher cortical levels. Some units exhibited significant nonlinear interactions between particular combinations of pitch, timbre, and azimuth. These interactions were most pronounced for pitch and timbre and were less commonly observed between spatial and nonspatial attributes. Such nonlinearities were most prevalent in primary auditory cortex, although they tended to be small compared with stimulus main effects.

Adult-like pitch processing skills in newborn infants

Event Abstract Back to Event Adult-like pitch processing skills in newborn infants G. Stefanics1, 2*, Gabor P. Háden1, 3, I. Sziller4, A. Beke4, M. D. Westergaard5, S. Denham6, L. Balázs1 and István Winkler1, 7 1 Institute for Psychology, Hungarian Academy of Sciences, Hungary 2 Department of Experimental Zoology and Neurobiology, University of Pécs, Hungary 3 Department of Cognitive Science, Budapest University of Technology and Economics, Hungary 4 First Department of Obstetrics and Gynecology, Semmelweis University, Hungary 5 Department of Physiology, Development and Neuroscience, University of Cambridge, United Kingdom 6 Centre for Theoretical and Computational Neuroscience, University of Plymouth, United Kingdom 7 Institute of Psychology, University of Szeged, Hungary Pitch is a perceived auditory feature ordering sounds on a scale from low to high. The auditory system analyzes sound structure in a complex way to arrive at perceiving and representing pitch. Some theories suggest that learning plays an important role in pitch perception. Therefore we tested whether or not nontrivial forms of pitch analysis are functional in newborn infants, especially because pitch processing is an important prerequisite of music and speech perception and is also crucial for representing auditory objects. In two separate studies we investigated 1) whether or not newborn infants generalize pitch across different timbres and 2) whether the auditory system of newborns extracts the constancy of a pitch interval from exemplars varying in absolute pitch. In both experiments we recorded event-related brain potentials in healthy newborn infants to frequent standard and infrequent deviant sound stimuli. In study 1) timbre-independent pitch representation was tested by a paradigm where we applied randomly varying tones of different perceived resonator size, which is an aspect of timbre that informs the listener about the size of the sound source. The elicitation of an early negative and a later positive discriminative response by deviant sounds demonstrated that the neonate auditory system represents pitch separately from timbre. In study 2) we investigated whether the auditory system of newborns extracts the constancy of a pitch interval from exemplars varying in absolute pitch. To this end we applied an oddball paradigm consisting of frequent standard and infrequent deviant tone pairs. Tone pairs varied in absolute frequency. Standard and deviant pairs differed in the amount of pitch difference within the pairs, but not in the direction of pitch change. Deviant tone pairs elicited a discriminative ERP response, suggesting that the neonate auditory system represents pitch intervals similarly to adults. Our results demonstrate that adult-like pitch processing abilities are present already at birth and they allow newborn infants to learn music, speech prosody, and to process various important auditory cues based on spectral acoustic features. Conference: MMN 09 Fifth Conference on Mismatch Negativity (MMN) and its Clinical and Scientific Applications, Budapest, Hungary, 4 Apr - 7 Apr, 2009. Presentation Type: Poster Presentation Topic: Poster Presentations Citation: Stefanics G, Háden GP, Sziller I, Beke A, Westergaard MD, Denham S, Balázs L and Winkler I (2009). Adult-like pitch processing skills in newborn infants. Conference Abstract: MMN 09 Fifth Conference on Mismatch Negativity (MMN) and its Clinical and Scientific Applications. doi: 10.3389/conf.neuro.09.2009.05.070 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 24 Mar 2009; Published Online: 24 Mar 2009. * Correspondence: G. Stefanics, Institute for Psychology, Hungarian Academy of Sciences, Budapest, Hungary, gstefanics@gmail.com Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers G. Stefanics Gabor P Háden I. Sziller A. Beke M. D Westergaard S. Denham L. Balázs István Winkler Google G. Stefanics Gabor P Háden I. Sziller A. Beke M. D Westergaard S. Denham L. Balázs István Winkler Google Scholar G. Stefanics Gabor P Háden I. Sziller A. Beke M. D Westergaard S. Denham L. Balázs István Winkler PubMed G. Stefanics Gabor P Háden I. Sziller A. Beke M. D Westergaard S. Denham L. Balázs István Winkler Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

Development of pitch representations in infancy

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Ambiguous Pitch and the Temporal Representation of Inharmonic Iterated Rippled Noise in the Ventral Cochlear Nucleus

Neural coding of the pitch of complex sounds is vital for animals' ability to communicate and to perceptually organize natural acoustic scenes. Harmonic complex sounds typically have a well defined pitch corresponding to their fundamental frequency, whereas inharmonic sounds can exhibit pitch ambiguity: their pitch can have more than one value. Iterated rippled noise (IRN), a common "pitch stimulus," is generated from broadband noise by a cascade of delay-and-add steps, with the delayed noise phase-shifted by varphi degrees. By varying varphi, the (in)harmonicity, and therefore the pitch ambiguity, of IRN can be manipulated. Recordings were made from single-units in the ventral cochlear nucleus of anesthetized guinea pigs in response to IRN and complex tones, systematically varying the inharmonicity. In their all-order interspike interval distributions, primary-like and chopper units tuned within the phase-locking range of best frequencies represent the waveform temporal fine structure (which varies with varphi). In contrast, those units tuned to higher frequencies represent the temporal-envelope modulation (independent of varphi). We show a temporal representation of ambiguous pitch for IRN and complex tones based on responses to the stimulus fine structure. Within the dominance region for pitch this representation follows the predictions of classic human behavioral experiments and provides a unifying contribution to possible neuro-temporal explanations for the pitch shift and pitch ambiguity associated with many inharmonic sounds.