Distinct Representations of Tonotopy and Pitch in Human Auditory Cortex.

Frequency-to-place mapping, or tonotopy, is a fundamental organizing principle throughout the auditory system, from the earliest stages of auditory processing in the cochlea to subcortical and cortical regions. Although cortical maps are referred to as tonotopic, it is unclear whether they simply reflect a mapping of physical frequency inherited from the cochlea, a computation of pitch based on the fundamental frequency, or a mixture of these two features. We used high-resolution functional magnetic resonance imaging (fMRI) to measure BOLD responses as male and female human participants listened to pure tones that varied in frequency or complex tones that varied in either spectral content (brightness) or fundamental frequency (pitch). Our results reveal evidence for pitch tuning in bilateral regions that partially overlap with the traditional tonotopic maps of spectral content. In general, primary regions within Heschl's gyri (HGs) exhibited more tuning to spectral content, whereas areas surrounding HGs exhibited more tuning to pitch.SIGNIFICANCE STATEMENT Tonotopy, an orderly mapping of frequency, is observed throughout the auditory system. However, it is not known whether the tonotopy observed in the cortex simply reflects the frequency spectrum (as in the ear) or instead represents the higher-level feature of fundamental frequency, or pitch. Using carefully controlled stimuli and high-resolution functional magnetic resonance imaging (fMRI), we separated these features to study their cortical representations. Our results suggest that tonotopy in primary cortical regions is driven predominantly by frequency, but also reveal evidence for tuning to pitch in regions that partially overlap with the tonotopic gradients but extend into nonprimary cortical areas. In addition to resolving ambiguities surrounding cortical tonotopy, our findings provide evidence that selectivity for pitch is distributed bilaterally throughout auditory cortex.

Cortical Correlates of Attention to Auditory Features.

Pitch and timbre are two primary features of auditory perception that are generally considered independent. However, an increase in pitch (produced by a change in fundamental frequency) can be confused with an increase in brightness (an attribute of timbre related to spectral centroid) and vice versa. Previous work indicates that pitch and timbre are processed in overlapping regions of the auditory cortex, but are separable to some extent via multivoxel pattern analysis. Here, we tested whether attention to one or other feature increases the spatial separation of their cortical representations and if attention can enhance the cortical representation of these features in the absence of any physical change in the stimulus. Ten human subjects (four female, six male) listened to pairs of tone triplets varying in pitch, timbre, or both and judged which tone triplet had the higher pitch or brighter timbre. Variations in each feature engaged common auditory regions with no clear distinctions at a univariate level. Attending to one did not improve the separability of the neural representations of pitch and timbre at the univariate level. At the multivariate level, the classifier performed above chance in distinguishing between conditions in which pitch or timbre was discriminated. The results confirm that the computations underlying pitch and timbre perception are subserved by strongly overlapping cortical regions, but reveal that attention to one or other feature leads to distinguishable activation patterns even in the absence of physical differences in the stimuli.SIGNIFICANCE STATEMENT Although pitch and timbre are generally thought of as independent auditory features of a sound, pitch height and timbral brightness can be confused for one another. This study shows that pitch and timbre variations are represented in overlapping regions of auditory cortex, but that they produce distinguishable patterns of activation. Most importantly, the patterns of activation can be distinguished based on whether subjects attended to pitch or timbre even when the stimuli remained physically identical. The results therefore show that variations in pitch and timbre are represented by overlapping neural networks, but that attention to different features of the same sound can lead to distinguishable patterns of activation.

Representations of Pitch and Timbre Variation in Human Auditory Cortex.

Pitch and timbre are two primary dimensions of auditory perception, but how they are represented in the human brain remains a matter of contention. Some animal studies of auditory cortical processing have suggested modular processing, with different brain regions preferentially coding for pitch or timbre, whereas other studies have suggested a distributed code for different attributes across the same population of neurons. This study tested whether variations in pitch and timbre elicit activity in distinct regions of the human temporal lobes. Listeners were presented with sequences of sounds that varied in either fundamental frequency (eliciting changes in pitch) or spectral centroid (eliciting changes in brightness, an important attribute of timbre), with the degree of pitch or timbre variation in each sequence parametrically manipulated. The BOLD responses from auditory cortex increased with increasing sequence variance along each perceptual dimension. The spatial extent, region, and laterality of the cortical regions most responsive to variations in pitch or timbre at the univariate level of analysis were largely overlapping. However, patterns of activation in response to pitch or timbre variations were discriminable in most subjects at an individual level using multivoxel pattern analysis, suggesting a distributed coding of the two dimensions bilaterally in human auditory cortex. SIGNIFICANCE STATEMENT: Pitch and timbre are two crucial aspects of auditory perception. Pitch governs our perception of musical melodies and harmonies, and conveys both prosodic and (in tone languages) lexical information in speech. Brightness-an aspect of timbre or sound quality-allows us to distinguish different musical instruments and speech sounds. Frequency-mapping studies have revealed tonotopic organization in primary auditory cortex, but the use of pure tones or noise bands has precluded the possibility of dissociating pitch from brightness. Our results suggest a distributed code, with no clear anatomical distinctions between auditory cortical regions responsive to changes in either pitch or timbre, but also reveal a population code that can differentiate between changes in either dimension within the same cortical regions.

Symmetric interactions and interference between pitch and timbre.

Variations in the spectral shape of harmonic tone complexes are perceived as timbre changes and can lead to poorer fundamental frequency (F0) or pitch discrimination. Less is known about the effects of F0 variations on spectral shape discrimination. The aims of the study were to determine whether the interactions between pitch and timbre are symmetric, and to test whether musical training affects listeners' ability to ignore variations in irrelevant perceptual dimensions. Difference limens (DLs) for F0 were measured with and without random, concurrent, variations in spectral centroid, and vice versa. Additionally, sensitivity was measured as the target parameter and the interfering parameter varied by the same amount, in terms of individual DLs. Results showed significant and similar interference between pitch (F0) and timbre (spectral centroid) dimensions, with upward spectral motion often confused for upward F0 motion, and vice versa. Musicians had better F0DLs than non-musicians on average, but similar spectral centroid DLs. Both groups showed similar interference effects, in terms of decreased sensitivity, in both dimensions. Results reveal symmetry in the interference effects between pitch and timbre, once differences in sensitivity between dimensions and subjects are controlled. Musical training does not reliably help to overcome these effects.