Behavioral genetics. Study suggests pitch perception is inherited.
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1. The distribution of binaural responses within the ferret's primary auditory cortex was determined by standard microelectrode mapping techniques. Single and multiple unit responses were recorded from the middle ectosylvian gyrus of barbiturate-anesthetized animals with tungsten microelectrodes (1.2-1.8 M omega) inserted into the brain perpendicular to the cortical surface. The characteristic frequency (CF) and binaural response characteristics were determined for each point sampled. 2. Gated pure tones were delivered independently to the two ears through a sealed sound system, and binaural response types were determined by comparison of evoked activity for monaural and binaural stimulation. Most binaural responses fell into one of three major groups: binaural summation (EE/F), binaural suppression (EO/I), or mixed binaural summation and suppression. 3. The neurons tended to be grouped together on the basis of their binaural response properties. Zones of binaural summation and suppression extended across the surface of the middle ectosylvian gyrus and intersected with regions of sound frequency representation. 4. Particular attention was paid to the distribution of binaural responses within isofrequency contours in the ferret's primary auditory cortex (AI). Along the length of each isofrequency contour, areas of EE/F alternated with areas of EO/I. The summation areas were typically between 0.5 and 0.7 mm wide. Single neurons with mixed binaural response properties were frequently found between groups of EE/F and EO/I cells. The mixed responses appeared to mark a transition in location between zones of summation and suppression responses. 5. The distribution of interaural intensity difference (IID) thresholds was also examined along the length of isofrequency contours. No systematic relation was found between IID threshold and the distance along an isofrequency contour.
In music, melodic information is thought to be encoded in two forms, a contour code (up/down pattern of pitch changes) and an interval code (pitch distances between successive notes). A recent study recording the mismatch negativity (MMN) evoked by pitch contour and interval deviations in simple melodies demonstrated that people with no formal music education process both contour and interval information in the auditory cortex automatically. However, it is still unclear whether musical experience enhances both strategies of melodic encoding. We designed stimuli to examine contour and interval information separately. In the contour condition there were eight different standard melodies (presented on 80% of trials), each consisting of five notes all ascending in pitch, and the corresponding deviant melodies (20%) were altered to descending on their final note. The interval condition used one five-note standard melody transposed to eight keys from trial to trial, and on deviant trials the last note was raised by one whole tone without changing the pitch contour. There was also a control condition, in which a standard tone (990.7 Hz) and a deviant tone (1111.0 Hz) were presented. The magnetic counterpart of the MMN (MMNm) from musicians and nonmusicians was obtained as the difference between the dipole moment in response to the standard and deviant trials recorded by magnetoencephalography. Significantly larger MMNm was present in musicians in both contour and interval conditions than in nonmusicians, whereas MMNm in the control condition was similar for both groups. The interval MMNm was larger than the contour MMNm in musicians. No hemispheric difference was found in either group. The results suggest that musical training enhances the ability to automatically register abstract changes in the relative pitch structure of melodies.
P300 evoked potential patterns were evaluated for two active and two passive attentional states in which stimuli were presented in either one or two sensory modalities. Latency to an infrequent auditory tone was shortest when counting the frequent stimulus, longer when counting the infrequent stimulus, longer still under a passive condition, and longest in a non-P300 condition. P300 amplitudes identifiable in the passive attentional state were enhanced when stimuli were from two, rather than from one, sensory modality. Implications for understanding the mechanism of attention are discussed. Preliminary findings in traumatic brain injury patients suggest that passive P300 responses may be useful in assessing the extent and severity of brain dysfunction.
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The Grammar of Dutch Intonation (GDI) provides a description of the possible intonation contours of Dutch. The GDI distinguishes accent-lending and nonaccent-lending pitch configurations, but refrains from further functional statements. This paper describes an experimental attempt to verify meaning hypotheses for four Dutch single-accent pitch patterns as postulated in the linguistic literature. The four pitch accent types were realized on proper names; the abstract meanings, in terms of the manipulation of an element of the background shared between speaker and listener, were incorporated in situational contexts, distinguishing between a "default" and a vocative use of the proper name ("orientation"). Listeners ranked the four melodic shapes from most to least appropriate in their specific context. After revision of part of the materials a second perception experiment was conducted, in which subjects had a rank four contexts from most to least appropriate for a specific pitch accent type. Results show a distinct effect of "orientation" on the appropriateness of two of the investigated pitch accent types in the various context types; the other two pitch accent types are associated with the predicted context types (and vice versa) well above chance, indicating the viability of at least two of the linguistic proposals.
The issue of the perception of lexical tone has been addressed mainly through studies of Southeast Asian languages which feature phonological contour tones as well as level tones. Little attention has been paid to African languages which have, almost exclusively, only level tones. This paper examines tone perception in Mambila, a Benue-Congo language with four level lexical tones. A categorization experiment was run to determine some of the salient aspects of the perceptual nature of these tones. Since the four tones are well defined with respect to production, we sought to determine whether this characteristic carried over into perception, the expectation being that experimental stimuli, on the basis of pitch height alone, would fall into four reasonably well defined categories. Results showed interesting differences across the four tones, with indications that the two Mid tones, T2 and T3, are perceptually different than the High (T1) and Low (T4) tones. The experiment was run a second time, using a group of native English listeners, to assess to what extent results for the Mambila listeners were determined by the perceptual structure of the Mambila tone system. A Signal Detection analysis was used, which revealed important differences between the two groups of listeners. Results are discussed in light of what is known about universal tendencies of tone systems and the historical development of the Mambila system.
Studies are reviewed from the perspective of a neurologist and epileptologist interested in "music and the brain." At the neurocognitive level, deficits in pitch discrimination of patients with brain lesions and those during the intracarotid amobarbital test are outlined, because they show that the temporal lobe and, in particular, the right acoustic cortex are crucial. Hallucinations of music during epileptic seizures as well as the analysis of musicogenic epilepsy point to the same gross localization and lateralization. At the esthetic level, music theoretical concepts on the consonance-dissonance dichotomy and related EEG examinations are reported, which illustrate the importance of mesiolimbic temporal lobe structures for the pleasure that we might experience when listening to music. The complex interaction of many neuronal circuits and assemblies of both hemispheres in musical perception and performance is illustrated by musical analysis of a recording by an organ player who experienced a right temporal lobe seizure. This analysis revealed that the seizure-induced errors of the left hand were compensated with the right hand in a musically meaningful way.
Two new empirical studies address the relationship between music and language. The first focuses on melody and uses research in phonetics to investigate the long-held notion that instrumental music reflects speech patterns in a composer's native language. The second focuses on syntax and addresses the relationship between musical and linguistic syntactic processing via the study of aphasia, an approach that has been explored very little. The results of these two studies add to a growing body of evidence linking music and language with regard to structural patterns and brain processing.
One approach to comparing the neural bases of language and music is through the use of song, which is a unique and ecological combination of these two cognitive domains. In song, language and music are merged into one acoustic signal with two salient dimensions. By manipulating either the linguistic or musical dimensions (or both) of song and studying their relationships, it is possible to gain important information about the neural networks underlying language and music cognition. We will present a brief review followed by recent behavioral, electrophysiological, and neuroimaging studies concerned with the functional and structural relationships of music and language. These results, together with the previous studies in the field, help understanding whether the different levels of music and language processing are independent or interactive.
By mere exposure to musical pieces in everyday life, Western listeners acquire sensitivity to the regularities of the tonal system and to the context dependency of musical sounds. This implicitly acquired tonal knowledge allows nonmusician listeners to perceive relationships among musical events and to develop expectations for future events that then influence the processing of these events. The musical priming paradigm is one method of the indirect investigation of listeners' tonal knowledge. It investigates the influence of a preceding context (with its musical structures and relationships) on the processing of a musical target event, without asking participants for direct evaluations. Behavioral priming data have provided evidence for facilitated processing of musically related events in comparison to unrelated and less-related events. The sensitivity of implicit investigations is further shown by I.R., a patient with severe amusia, showing spared implicit knowledge of music. Finally, the priming paradigm allows us to investigate the neural correlates of musical structure processing. Two fMRI studies reported the implication of inferior frontal regions in musical priming, contrasting related and unrelated events, as well as finer structural manipulations contrasting in-key events.
As the functional neuroimaging literature grows, it becomes increasingly apparent that music and musical activities engage diverse regions of the brain. In this paper I discuss two studies to illustrate that exactly which brain areas are observed to be responsive to musical stimuli and tasks depends on the tasks and the methods used to describe the tasks and the stimuli. In one study, subjects listened to polyphonic music and were asked to either orient their attention selectively to individual instruments or in a divided or holistic manner across multiple instruments. The network of brain areas that was recruited changed subtly with changes in the task instructions. The focus of the second study was to identify brain regions that follow the pattern of movement of a continuous melody through the tonal space defined by the major and minor keys of Western tonal music. Such an area was identified in the rostral medial prefrontal cortex. This observation is discussed in the context of other neuroimaging studies that implicate this region in inwardly directed mental states involving decisions about the self, autobiographical memory, the cognitive regulation of emotion, affective responses to musical stimuli, and familiarity judgments about musical stimuli. Together with observations that these regions are among the last to atrophy in Alzheimer disease, and that these patients appear to remain responsive to autobiographically salient musical stimuli, very early evidence is emerging from the literature for the hypothesis that the rostral medial prefrontal cortex is a node that is important for binding music with memories within a broader music-responsive network.
Although considerable progress has been made recently in our understanding of the coding of complex sounds in the cerebral cortex, the processing and storage of tone sequences is still poorly understood. We have used functional magnetic resonance imaging to identify brain mechanisms involved in the encoding and retrieval of melodies by studying the anticipation of familiar music. The results suggest a specific role for each of the following brain structures: the anterior part of the right superior temporal cortex, the right inferior frontal cortex and anterior insula, the left anterior prefrontal cortex, the lateral cerebellum, and the anterior cingulate. In a separate study, we investigated single-neuron responses in the auditory cortex of awake behaving monkeys to alternating tone sequences that in humans evoke the perception of "streaming." Depending on the frequency separation between the tones, an initial single stream may segregate into two streams after a build-up period of several seconds. The neural responses in the monkeys' primary auditory cortex (A1) mirror the psychophysical time course extremely well, suggesting that habituation within A1 may be one reason for stream segregation. However, the higher auditory and prefrontal areas found to be activated by musical melodies are expected to interact with primary areas in both bottom-up and top-down fashion to bring about the perceptual organization of sound sequences.
This study was concerned with self-reported "tone deafness" and its possible relationship to congenital amusia. Nearly 17% of over 2,000 first-year psychology students at Queen's University self-reported tone deafness. Two hundred students were recruited from this pool of students, comprising 100 who reported tone deafness and 100 who reported that they were not tone-deaf (NTD). The study contained two parts. In part 1, participants completed the six tests of the Montreal Battery of Evaluation of Amusia (MBEA) developed by Peretz and collaborators. In part 2, participants completed an extensive questionnaire designed to elicit details about musical experiences, abilities, training, and interests. Twenty-eight questionnaire items allowing a quantitative response were subjected to factor analysis. Four orthogonal components emerged from the analysis. The components reflected self-report of (1) vocal production, (2) music instruction, (3) listening attitudes, and (4) childhood memories of musical environment. Results for each of the MBEA tests and composite scores for all tests were regressed on participants' factor scores. The best and significant predictors of the MBEA scores were factor I and factor II, followed by factor III. Factor scores accounted for a higher percentage of the variance in MBEA composite test results (27%) than the self-report of tone deafness alone (7%). The musical difficulties revealed by the MBEA test results for some participants warrant further attention and study. However, an encouraging conclusion from the MBEA results is that many individuals who consider themselves "tone-deaf" may not, in fact, have perceptual difficulties, and these individuals should be supported in any of their efforts to proceed with music enjoyment and instruction.
The musical pitch of harmonic complex sounds, such as instrumental sounds, is perceived primarily by decoding either the fundamental pitch (keynote) or spectral aspects of the stimuli, for example, single harmonics. We divided 334 professional musicians, including symphony orchestra musicians, 75 amateur musicians, and 54 nonmusicians, into either fundamental pitch listeners or spectral pitch listeners. We observed a strong correlation between pitch perception preference and asymmetry of brain structure and function in the pitch-sensitive lateral areas of Heschl's gyrus (HG), irrespective of musical ability. In particular, fundamental pitch listeners exhibited both larger gray matter volume measured using magnetic resonance imaging (MRI) and enhanced P50m activity measured using magnetoencephalography (MEG) in the left lateral HG, which is sensitive to rapid temporal processing. Their chosen instruments were percussive or high-pitched instruments that produce short, sharp, or impulsive tones (e.g., drums, guitar, piano, trumpet, or flute). By contrast, spectral pitch listeners exhibited a dominant right lateral HG, which is known to be sensitive to slower temporal and spectral processing. Their chosen instruments were lower-pitched melodic instruments that produce rather sustained tones with characteristic changes in timbre (e.g., bassoon, saxophone, french horn, violoncello, or organ). Singers also belonged to the spectral pitch listeners. Furthermore, the absolute size of the neural HG substrate depended strongly on musical ability. Overall, it is likely that both magnitude and asymmetry of lateral HG, and the related perceptual mode, may have an impact on preference for particular musical instruments and on musical performance.
Our results suggest that musical training alters the functional anatomy of rapid spectrotemporal processing, resulting in improved behavioral performance along with a more efficient functional network primarily involving traditional language regions. This finding may have important implications for improving language/reading skills, especially in children struggling with dyslexia.
We trained musically naive subjects to play a short piano melody by ear in a fully monitored computerized environment and tested their potential to acquire a functional linkage between actions and sounds. Individual notes that were simply acoustic pretraining signals became "physically meaningful" posttraining. In addition, we found preliminary evidence that passive listening to a newly learned musical piece can enhance motor performance in the absence of physical practice.
Temporal processing of periodic acoustic signals in the auditory brain stem provides an explanation for pitch perception and the natural preference of our hearing system for harmonic relationships in music. Experimental evidence is reviewed for a corresponding neuronal model of correlation analysis and the spatial representation of pitch information along the second neural axis of the auditory system.