Congenital amusia is currently thought to be a life-long neurogenetic disorder in music perception, impervious to training in pitch or melody discrimination. This study provides an explicit test of whether amusic deficits can be reduced with training. Twenty amusics and 20 matched controls participated in four sessions of psychophysical training involving either pure-tone (500 Hz) pitch discrimination or a control task of lateralization (interaural level differences for bandpass white noise). Pure-tone pitch discrimination at low, medium, and high frequencies (500, 2000, and 8000 Hz) was measured before and after training (pretest and posttest) to determine the specificity of learning. Melody discrimination was also assessed before and after training using the full Montreal Battery of Evaluation of Amusia, the most widely used standardized test to diagnose amusia. Amusics performed more poorly than controls in pitch but not localization discrimination, but both groups improved with practice on the trained stimuli. Learning was broad, occurring across all three frequencies and melody discrimination for all groups, including those who trained on the non-pitch control task. Following training, 11 of 20 amusics no longer met the global diagnostic criteria for amusia. A separate group of untrained controls (n = 20), who also completed melody discrimination and pretest, improved by an equal amount as trained controls on all measures, suggesting that the bulk of learning for the control group occurred very rapidly from the pretest. Thirty-one trained participants (13 amusics) returned one year later to assess long-term maintenance of pitch and melody discrimination. On average, there was no change in performance between posttest and one-year follow-up, demonstrating that improvements on pitch- and melody-related tasks in amusics and controls can be maintained. The findings indicate that amusia is not always a life-long deficit when using the current standard diagnostic criteria.

Pitch perception and pitch matching may link to individual reading skills.

Vocal pitch discrimination abilities were compared in sixteen children with vocal fold nodules (CwVN) and sixteen matched controls with typical voices (CwTV). Vocal pitch discrimination was also evaluated in thirty-five vocally healthy children and twenty adults to examine potential changes as a function of maturation. CwTV were categorized as either younger (N = 15, 5.6-7.7 years) or older (N = 20, 8.2-11.7 years). Participants completed two-alternative, forced choice listening tasks in which they judged whether pairs of sustained /α/ tokens were different in pitch. Each pair consisted of a base token with a fundamental frequency fo ) of 216.2 Hz and a test token with a fo that was adaptively modified, according to the participant's prior judgments. There were no significant differences in pitch discrimination abilities between CwVN and CwTV. Pitch discrimination abilities were significantly poorer in younger and older CwTV as compared to adults. Additionally, younger CwTV had significantly poorer discrimination abilities than older CwTV. Findings from this study suggest that CwVN do not have differences in pitch discrimination abilities, yet, therapies designed for CwVN should consider this developmental trend in perceptual abilities.

The cochlear implant (CI) is the most widely used neuroprosthesis, recovering hearing for more than half a million severely-to-profoundly hearing-impaired people. However, CIs still have significant limitations, with users having severely impaired pitch perception. Pitch is critical to speech understanding (particularly in noise), to separating different sounds in complex acoustic environments, and to music enjoyment. In recent decades, researchers have attempted to overcome shortcomings in CIs by improving implant technology and surgical techniques, but with limited success. In the current study, we take a new approach of providing missing pitch information through haptic stimulation on the forearm, using our new mosaicOne_B device. The mosaicOne_B extracts pitch information in real-time and presents it via 12 motors that are arranged in ascending pitch along the forearm, with each motor representing a different pitch. In normal-hearing subjects listening to CI simulated audio, we showed that participants were able to discriminate pitch differences at a similar performance level to that achieved by normal-hearing listeners. Furthermore, the device was shown to be highly robust to background noise. This enhanced pitch discrimination has the potential to significantly improve music perception, speech recognition, and speech prosody perception in CI users.

Previous studies indicate that temporal predictability can enhance timing and intensity perception, but it is not known whether it also enhances pitch perception, despite pitch being a fundamental perceptual attribute of sound. Here we investigate this in the context of rhythmic regularity, a form of predictable temporal structure common in sound streams, including music and speech. It is known that neural oscillations in low (delta: 1-3 Hz) and high (beta: 15-25 Hz) frequency bands entrain to rhythms in phase and power, respectively, but it is not clear why both low and high frequency bands entrain to external rhythms, and whether they and their coupling serve different perceptual functions. Participants discriminated near-threshold pitch deviations (targets) embedded in either rhythmic (regular/isochronous) or arrhythmic (irregular/non-isochronous) tone sequences. Psychophysically, we found superior pitch discrimination performance for target tones in rhythmic compared to arrhythmic sequences. Electroencephalography recordings from auditory cortex showed that delta phase, beta power modulation, and delta-beta coupling were all modulated by rhythmic regularity. Importantly, trial-by-trial neural-behavioural correlational analyses showed that, prior to a target, the depth of U-shaped beta power modulation predicted pitch discrimination sensitivity whereas cross-frequency coupling strength predicted reaction time. These novel findings suggest that delta phase might reflect rhythmic temporal expectation, beta power temporal attention, and delta-beta coupling auditory-motor communication. Together, low and high frequency auditory neural oscillations reflect different perceptual functions that work in concert for tracking rhythmic regularity and proactively facilitate pitch perception.

Growing neuroimaging and clinical evidence suggests that the cerebellum plays a critical role in perception. In the auditory domain, the cerebellum seems to be important in different aspects of music and sound processing. Here we investigated the possible causal role of the cerebellum in two auditory tasks, a pitch discrimination and a timbre discrimination task. Specifically, participants performed a pitch and a timbre discrimination task prior and after receiving offline low frequency transcranical magnetic stimulation (TMS) over their (right) cerebellum.

Harmonic complex tones are easier to detect in noise than inharmonic complex tones, providing a potential perceptual advantage in complex auditory environments. Here, we explored whether the harmonic advantage extends to other auditory tasks that are important for navigating a noisy auditory environment, such as amplitude- and frequency-modulation detection. Sixty young normal-hearing listeners were tested, divided into two equal groups with and without musical training. Consistent with earlier studies, harmonic tones were easier to detect in noise than inharmonic tones, with a signal-to-noise ratio (SNR) advantage of about 2.5 dB, and the pitch discrimination of the harmonic tones was more accurate than that of inharmonic tones, even after differences in audibility were accounted for. In contrast, neither amplitude- nor frequency-modulation detection was superior with harmonic tones once differences in audibility were accounted for. Musical training was associated with better performance only in pitch-discrimination and frequency-modulation-detection tasks. The results confirm a detection and pitch-perception advantage for harmonic tones but reveal that the harmonic benefits do not extend to suprathreshold tasks that do not rely on extracting the fundamental frequency. A general theory is proposed that may account for the effects of both noise and memory on pitch-discrimination differences between harmonic and inharmonic tones.

Behavioral adaption to a changing environment is critical for an animal's survival. How well the brain can modify its functional properties based on experience essentially defines the limits of behavioral adaptation. In adult animals the extent to which experience shapes brain function has not been fully explored. Moreover, the perceptual consequences of experience-induced changes in the brains of adults remain unknown. Here we show that the tonotopic map in the primary auditory cortex of adult rats living with low-level ambient noise underwent a dramatic reorganization. Behaviorally, chronic noise-exposure impaired fine, but not coarse pitch discrimination. When tested in a noisy environment, the noise-exposed rats performed as well as in a quiet environment whereas the control rats performed poorly. This suggests that noise-exposed animals had adapted to living in a noisy environment. Behavioral pattern analyses revealed that stress or distraction engendered by the noisy background could not account for the poor performance of the control rats in a noisy environment. A reorganized auditory map may therefore have served as the neural substrate for the consistent performance of the noise-exposed rats in a noisy environment.

Can human listeners use implicit temporal contingencies in auditory input to form temporal predictions, and if so, how are these predictions represented endogenously? To assess this question, we implicitly manipulated temporal predictability in an auditory pitch discrimination task: unbeknownst to participants, the pitch of the standard tone could either be deterministically predictive of the temporal onset of the target tone, or convey no predictive information. Predictive and non-predictive conditions were presented interleaved in one stream, and separated by variable inter-stimulus intervals such that there was no dominant stimulus rhythm throughout. Even though participants were unaware of the implicit temporal contingencies, pitch discrimination sensitivity (the slope of the psychometric function) increased when the onset of the target tone was predictable in time (N = 49, 28 female, 21 male). Concurrently recorded EEG data (N = 24) revealed that standard tones that conveyed temporal predictions evoked a more negative N1 component than non-predictive standards. We observed no significant differences in oscillatory power or phase coherence between conditions during the foreperiod. Importantly, the phase angle of delta oscillations (1-3 Hz) in auditory areas in the post-standard and pre-target time windows predicted behavioral pitch discrimination sensitivity. This suggests that temporal predictions are encoded in delta oscillatory phase during the foreperiod interval. In sum, we show that auditory perception benefits from implicit temporal contingencies, and provide evidence for a role of slow neural oscillations in the endogenous representation of temporal predictions, in absence of exogenously driven entrainment to rhythmic input.

We examined effects of significance of task irrelevant changes in the location of tones on the mismatch negativity (MMN) and P3a event related brain potentials. The participants were to discriminate between two frequency modulated tones differing from each other in the direction of frequency glide. Each tone was delivered through one of five loudspeakers in front of the participant. On most trials, a tone was presented from the same location as the preceding tone, but occasionally the location changed. In the Varying Location Condition, these changes, although irrelevant with regard to pitch discrimination, were still significant for performance as the following tones were presented from the new location where attention had to be therefore shifted. In the Fixed Location Condition, the location changes were less significant as the tones following a location change were presented from the original location. In both conditions, the location changes were associated with decreased hit rates and increased reaction times in the pitch discrimination task. However, the hit rate decrease was larger in the Fixed Location Condition suggesting that in this condition the location changes were just distractors. MMN and P3a responses were elicited by location changes in both conditions. In the Fixed Location Condition, a P3a was also elicited by the first tone following a location change at the original location while the MMN was not. Thus, the P3a appeared to be related to shifting of attention in space and was not tightly coupled with MMN elicitation.

Accurate pitch perception of harmonic complex tones is widely believed to rely on temporal fine structure information conveyed by the precise phase-locked responses of auditory-nerve fibers. However, accurate pitch perception remains possible even when spectrally resolved harmonics are presented at frequencies beyond the putative limits of neural phase locking, and it is unclear whether residual temporal information, or a coarser rate-place code, underlies this ability. We addressed this question by measuring human pitch discrimination at low and high frequencies for harmonic complex tones, presented either in isolation or in the presence of concurrent complex-tone maskers. We found that concurrent complex-tone maskers impaired performance at both low and high frequencies, although the impairment introduced by adding maskers at high frequencies relative to low frequencies differed between the tested masker types. We then combined simulated auditory-nerve responses to our stimuli with ideal-observer analysis to quantify the extent to which performance was limited by peripheral factors. We found that the worsening of both frequency discrimination and F0 discrimination at high frequencies could be well accounted for (in relative terms) by optimal decoding of all available information at the level of the auditory nerve. A Python package is provided to reproduce these results, and to simulate responses to acoustic stimuli from the three previously published models of the human auditory nerve used in our analyses.

Subcortical auditory nuclei contribute to pitch perception, but how subcortical sound encoding is related to pitch processing for music perception remains unclear. Conventionally, enhanced subcortical sound encoding is considered underlying superior pitch discrimination. However, associations between superior auditory perception and the context-dependent plasticity of subcortical sound encoding are also documented. Here, we explored the subcortical neural correlates to music pitch perception by analyzing frequency-following responses (FFRs) to musical sounds presented in a predictable context and a random context. We found that the FFR inter-trial phase-locking (ITPL) was negatively correlated with behavioral performances of discrimination of pitches in music melodies. It was also negatively correlated with the plasticity indices measuring the variability of FFRs to physically identical sounds between the two contexts. The plasticity indices were consistently positively correlated with pitch discrimination performances, suggesting the subcortical context-dependent plasticity underlying music pitch perception. Moreover, the raw FFR spectral strength was not significantly correlated with pitch discrimination performances. However, it was positively correlated with behavioral performances when the FFR ITPL was controlled by partial correlations, suggesting that the strength of subcortical sound encoding underlies music pitch perception. When the spectral strength was controlled by partial correlations, the negative ITPL-behavioral correlations were maintained. Furthermore, the FFR ITPL, the plasticity indices, and the FFR spectral strength were more correlated with pitch than with rhythm discrimination performances. These findings suggest that the context-dependent plasticity and the strength of subcortical encoding of musical sounds are independently and perhaps specifically associated with pitch perception for music melodies.

Aptitude for and proficiency in acquiring new languages varies in the human population but their neural bases are largely unknown. We investigated the influence of cortical thickness on language learning predictors measured by the LLAMA tests and a pitch-change discrimination test. The LLAMA tests are first language-independent assessments of language learning aptitude for vocabulary, phonetic working memory, sound-symbol correspondence (not used in this study), and grammatical inferencing. Pitch perception proficiency is known to predict aptitude for learning new phonology. Results show a correlation between scores in a grammatical meaning-inferencing aptitude test and cortical thickness of Broca's area (r(30) = 0.65, p = 0.0202) and other frontal areas (r(30) = 0.66, p = 0.0137). Further, a correlation was found between proficiency in discriminating pitch-change direction and cortical thickness of the right Broca homologue (r(30) = 0.57, p = 0.0006). However, no correlations were found for aptitude for vocabulary learning or phonetic working memory. Results contribute to locating cortical regions important for language-learning aptitude.

Musical training has been shown to have a positive influence on a variety of skills, including auditory-based tasks and nonmusical cognitive and executive functioning tasks; however, because previous investigations have yielded mixed results regarding the relationship between musical training and these skills, the purpose of this study was to examine and compare the auditory processing skills of children who receive focused, daily musical training with those with more limited, generalized musical training. Sixteen typically developing children (second-fourth grade) from two different schools receiving different music curricula were assessed on measures of pitch discrimination, temporal sequencing, and prosodic awareness. The results indicated significantly better scores in pitch discrimination abilities for the children receiving daily, focused musical training (School 1) compared to students attending music class only once per week, utilizing a more generalized elementary school music curriculum (School 2). The findings suggest that more in-depth and frequent musical training may be associated with better pitch discrimination abilities in children. This finding is important given that the ability to discriminate pitch has been linked to improved phonological processing skills, an important skill for developing spoken language and literacy. Future investigations are needed to determine whether the null findings for temporal sequencing and prosodic awareness can be replicated or may be different for various grades and tasks for measuring these abilities.

Pitch accents are local pitch patterns that convey differences in word prominence and modulate the information structure of the discourse. Despite the importance to discourse in languages like English, neural processing of pitch accents remains understudied. The current study investigates the neural processing of pitch accents by native and non-native English speakers while they are listening to or ignoring 45 min of continuous, natural speech. Leveraging an approach used to study phonemes in natural speech, we analyzed thousands of electroencephalography (EEG) segments time-locked to pitch accents in a prosodic transcription. The optimal neural discrimination between pitch accent categories emerged at latencies between 100 and 200 ms. During these latencies, we found a strong structural alignment between neural and phonetic representations of pitch accent categories. In the same latencies, native listeners exhibited more robust processing of pitch accent contrasts than non-native listeners. However, these group differences attenuated when the speech signal was ignored. We can reliably capture the neural processing of discrete and contrastive pitch accent categories in continuous speech. Our analytic approach also captures how language-specific knowledge and selective attention influences the neural processing of pitch accent categories.

Background: Perceptual and electrophysiological studies have found reduced speech discrimination in quiet and noisy environment, delayed neural timing, decreased neural synchrony, and decreased temporal processing ability in elderlies, even those with normal hearing. However, recent studies have also demonstrated that language experience and auditory training enhance the temporal dynamics of sound encoding in the auditory brainstem response (ABR). The purpose of this study was to explore the pitch processing ability at the brainstem level in an aging population that has a tonal language background. Method: Mandarin speaking younger (n = 12) and older (n = 12) adults were recruited for this study. All participants had normal audiometric test results and normal suprathreshold click-evoked ABR. To record frequency following responses (FFRs) elicited by Mandarin lexical tones, two Mandarin Chinese syllables with different fundamental frequency pitch contours (Flat Tone and Falling Tone) were presented at 70 dB SPL. Fundamental frequencies (f0) of both the stimulus and the responses were extracted and compared to individual brainstem responses. Two indices were used to examine different aspects of pitch processing ability at the brainstem level: Pitch Strength and Pitch Correlation. Results: Lexical tone elicited FFR were overall weaker in the older adult group compared to their younger adult counterpart. Measured by Pitch Strength and Pitch Correlation, statistically significant group differences were only found when the tone with a falling f0 (Falling Tone) were used as the stimulus. Conclusion: Results of this study demonstrated that in a tonal language speaking population, pitch processing ability at the brainstem level of older adults are not as strong and robust as their younger counterparts. Findings of this study are consistent with previous reports on brainstem responses of older adults whose native language is English. On the other hand, lexical tone elicited FFRs have been shown to correlate with the length of language exposure. Older adults' degraded responses in our study may also be due to that, the Mandarin speaking older adults' long term exposure somewhat counteracted the negative impact on aging and helped maintain, or at least reduced, the degradation rate in their temporal processing capacity at the brainstem level.

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.

THIS FUNCTIONAL MAGNETIC RESONANCE IMAGING STUDY EXAMINES SHARED AND DISTINCT CORTICAL AREAS INVOLVED IN THE AUDITORY PERCEPTION OF SONG AND SPEECH AT THE LEVEL OF THEIR UNDERLYING CONSTITUENTS: words and pitch patterns. Univariate and multivariate analyses were performed to isolate the neural correlates of the word- and pitch-based discrimination between song and speech, corrected for rhythmic differences in both. Therefore, six conditions, arranged in a subtractive hierarchy were created: sung sentences including words, pitch and rhythm; hummed speech prosody and song melody containing only pitch patterns and rhythm; and as a control the pure musical or speech rhythm. Systematic contrasts between these balanced conditions following their hierarchical organization showed a great overlap between song and speech at all levels in the bilateral temporal lobe, but suggested a differential role of the inferior frontal gyrus (IFG) and intraparietal sulcus (IPS) in processing song and speech. While the left IFG coded for spoken words and showed predominance over the right IFG in prosodic pitch processing, an opposite lateralization was found for pitch in song. The IPS showed sensitivity to discrete pitch relations in song as opposed to the gliding pitch in speech. Finally, the superior temporal gyrus and premotor cortex coded for general differences between words and pitch patterns, irrespective of whether they were sung or spoken. Thus, song and speech share many features which are reflected in a fundamental similarity of brain areas involved in their perception. However, fine-grained acoustic differences on word and pitch level are reflected in the IPS and the lateralized activity of the IFG.

Cochlear implant users hear pitch evoked by stimulation rate, but discrimination diminishes for rates above 300 Hz. This upper limit on rate pitch is surprising given the remarkable and specialized ability of the auditory nerve to respond synchronously to stimulation rates at least as high as 3 kHz and arguably as high as 10 kHz. Sensitivity to stimulation rate as a pitch cue varies widely across cochlear implant users and can be improved with training. The present study examines individual differences and perceptual learning of stimulation rate as a cue for pitch ranking. Adult cochlear implant users participated in electrode psychophysics that involved testing once per week for three weeks. Stimulation pulse rate discrimination was measured in bipolar and monopolar configurations for apical and basal electrodes. Base stimulation rates between 100 and 800 Hz were examined. Individual differences were quantified using psychophysically derived metrics of spatial tuning and temporal integration. This study examined distribution of measures across subjects, predictive power of psychophysically derived metrics of spatial tuning and temporal integration, and the effect of training on rate discrimination thresholds. Psychophysical metrics of spatial tuning and temporal integration were not predictive of stimulation rate discrimination, but discrimination thresholds improved at lower frequencies with training. Since most clinical devices do not use variable stimulation rates, it is unknown to what extent recipients may learn to use stimulation rate cues if provided in a clear and consistent manner.

Although aging-related alterations in the auditory sensory memory and involuntary change discrimination have been widely studied, it remains controversial whether the mismatch negativity (MMN) or its magnetic counterpart (MMNm) is modulated by physiological aging. This study aimed to examine the effects of aging on mismatch activity to pitch deviants by using a whole-head magnetoencephalography (MEG) together with distributed source modeling analysis. The neuromagnetic responses to oddball paradigms consisting of standards (1000 Hz, p=0.85) and deviants (1100 Hz, p=0.15) were recorded in healthy young (n=20) and aged (n=18) male adults. We used minimum norm estimate of source reconstruction to characterize the spatiotemporal neural dynamics of MMNm responses. Distributed activations to MMNm were identified in the bilateral fronto-temporo-parietal areas. Compared to younger participants, the elderly exhibited a significant reduction of cortical activation in bilateral superior temporal guri, superior temporal sulci, inferior fontal gyri, orbitofrontal cortices and right inferior parietal lobules. In conclusion, our results suggest an aging-related decline in auditory sensory memory and automatic change detection as indexed by MMNm.