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Music and the brain: disorders of musical listening.

The study of the brain bases for normal musical listening has advanced greatly in the last 30 years. The evidence from basic and clinical neuroscience suggests that listening to music involves many cognitive components with distinct brain substrates. Using patient cases reported in the literature, we develop an approach for understanding disordered musical listening that is based on the systematic assessment of the perceptual and cognitive analysis of music and its emotional effect. This approach can be applied both to acquired and congenital deficits of musical listening, and to aberrant listening in patients with musical hallucinations. Both the bases for normal musical listening and the clinical assessment of disorders now have a solid grounding in systems neuroscience.

Auditory Perception↗

Representation of concurrent acoustic objects in primary auditory cortex.

Auditory scene analysis involves the simultaneous grouping and parsing of acoustic data into separate mental representations (i.e., objects). Over two experiments, we examined the sequence of neural processes underlying concurrent sound segregation by means of recording of human middle latency auditory evoked responses. Participants were presented with complex sounds comprising several harmonics, one of which could be mistuned such that it was not an integer multiple of the fundamental frequency. In both experiments, Na (approximately 22 ms) and Pa (approximately 32 ms) waves were reliably generated for all classes of stimuli. For stimuli with a fundamental frequency of 200 Hz, the mean Pa amplitude was significantly larger when the third harmonic was mistuned by 16% of its original value, relative to when it was tuned. The enhanced Pa amplitude was related to an increased likelihood in reporting the presence of concurrent auditory objects. Our results are consistent with a low-level stage of auditory scene analysis in which acoustic properties such as mistuning act as preattentive segregation cues that can subsequently lead to the perception of multiple auditory objects.

Adult↗

Brain networks that track musical structure.

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.

Acoustic Stimulation↗

Frequency-specific auditory brainstem responses to clicks masked by notched noise.

Frequency-specific auditory brainstem responses were recorded as a function of the intensity of a click masked by notch-filtered white noise (i.e. band-stop-filtered). The bandwidth of the notch was fixed at five-thirds octave and the center frequency was 0.5, 1.0, 2.0 or 4.0 kHz. For normally hearing subjects the latency of the main peak in the responses is increasing with decreasing center frequency of the notch and with decreasing click intensity. At a click sensation level of 60 dB, the mean of 7 subjects' peak latency was 6.7 ms for the center frequency 4.0 kHz and 10.2 ms for the center frequency 0.5 kHz. The thresholds derived from the frequency-specific responses were within 20 dB above the subjective thresholds of audibility of the clicks as heard when masked by notched noise.

Adult↗

Auditory localization of nearby sources. Head-related transfer functions.

Although researchers have long recognized the unique properties of the head-related transfer function (HRTF) for nearby sources (within 1 m of the listener's head), virtually all of the HRTF measurements described in the literature have focused on source locations 1 m or farther from the listener. In this study, HRTFs for sources at distances from 0.12 to 1 m were calculated using a rigid-sphere model of the head and measured using a Knowles Electronic Manikin for Acoustic Research (KEMAR) and an acoustic point source. Both the calculations and the measurements indicate that the interaural level difference (ILD) increases substantially for lateral sources as distance decreases below 1 m, even at low frequencies where the ILD is small for distant sources. In contrast, the interaural time delay (ITD) is roughly independent of distance even when the source is close. The KEMAR measurements indicate that the direction of the source relative to the outer ear plays an important role in determining the high-frequency response of the HRTF in the horizontal plane. However, the elevation-dependent characteristics of the HRTFs are not strongly dependent on distance, and the contribution of the pinna to the HRTF is independent of distance beyond a few centimeters from the ear. Overall, the results suggest that binaural cues play an important role in auditory distance perception for nearby sources.

Dichotic Listening Tests↗

[Recent trends in the psychology of music perception: the perceptual processes of melody and musical pitch].

In this article some of the recent researches on music perception are reviewed, with a special emphasis upon those that concern melody organization and musical pitch. Lerdahl and Jackendoff (1983) pointed out that there is a grammatical parallel between music and language, and presented some grammatical rules for the tonal music. Serafine (1983) cautioned us not to confuse the "style principles" with the "musical cognitive process". The recent experiments on musical pitch and melody has reinforced the psychological validity of musical segmentation, tonality, pitch chroma, scale, and key. Theoretical approaches to the relations in the scale systems gave the possibility of a "new scale system" for use in psychological experiments, while the recent pitch models has made it possible for us to control tonality and to generate tones never used before in playing music. The need for a greater cross-stylistic generality of psychological concepts in music and the farther clarification of the "musical cognitive process" through experiments utilizing the "new scale system" and the "new tones" are indicated.

Auditory Perception↗

Responses from AVCN units in the cat before and after inducement of an acute noise trauma.

Acoustically evoked responses of single units in the anteroventral part of the cochlear nucleus (AVCN) in the cat were studied together with compound eighth-nerve action potentials (AP). Halfway through the experiments the cats were exposed for half an hour to pink noise at 105 dB SPL, producing an average threshold shift of 30 dB (maximum 50 dB) in the 2-6 kHz region. The effect of noise exposure was studied in two ways. On the one hand we compared the results for one population of units measured before the noise exposure with those found for another population measured afterwards. On the other hand we compared the results before and after the noise exposure for one unit that could be kept under observation during the noise exposure. After the noise exposure spontaneous activity and phase-locking of the responses of the units to the stimulus waveform were not significantly different from the pre-exposure findings. Response latency tended to increase. Therefore, the decrease of latency found for APs must be due to a shift to higher frequencies of the population of units contributing to the AP. The sharply tuned tip segments of tuning curves shift to higher levels whereas the low-frequency tails remain at about the same level. Q10 decreases by at most 50%, which was also found for AP tuning curves. Response spectra for clicks and noise (reverse correlation function) did not show a significant decrease of frequency selectivity. Units with CF less than 3 kHz may show a shift of CF to a lower frequency by 10-20%. After inducement of the noise trauma the sharply tuned tip segment of a tuning curve may not be found and CF may be assigned to a local minimum in the low-frequency tail of the tuning curve.

Animals↗

On the capability of the vestibular apparatus to perceive sound stimuli.

This paper discusses the capacity of sound perception in 5 profoundly deaf subjects in whom a "fenestra" was previously formated on the lateral semicircular canal. The ability of sound and vibration perception after fenestra formation in profoundly deaf subjects with normal function of the vestibular apparatus has been described previously. The experiments for this procedure were done on pigeons by Wit & Bleeker. The subjects aged from 22 to 34 years. Causes of deafness varied. Their hearing threshold was bilaterally higher than 95 dB HL and their vestibular apparatus was of normal excitability bilaterally when tested with a caloric test. In all 5 patients the ear conduction hearing threshold remained the same, but the ability of vibration perception improved significantly, and ranged postoperatively between 30 and 45 dB HL. The frequency dynamic range was from 125 to 2,000 Hz in 4 subjects, and from 125 to 4,000 Hz in 1 subject. The patients claimed to have perceived higher intensities, but with a variable dynamic range from frequency to frequency. The intensity range amounted to between 15 and 40 dB. The obtained data of vibration perception ability are discussed.

Adult↗

Characterization and simulation of impaired hearing: implications for hearing aid design.

This article reviews several aspects of auditory perception that are affected by hearing loss of cochlear origin. It is argued that most of the observed effects can be understood in terms of damage to a physiologically vulnerable active process in the cochlea. In a normal ear, this process enhances sensitivity and frequency selectivity and reduces the slope of the input-output function on the basilar membrane. Damage to the active process in impaired ears leads to reduced sensitivity and frequency selectivity, an abnormally rapid rate of growth of loudness with intensity, reduced temporal integration and, for certain types of stimuli, reduced temporal resolution. The implications and relevance of each of these effects to the design and fitting of hearing aids are discussed.

Acoustics↗

Predicting auditory tone-in-noise detection performance: the effects of neural variability.

Collecting and analyzing psychophysical data is a fundamental mechanism for the study of auditory processing. However, because this approach relies on human listening experiments, it can be costly in terms of time and money spent gathering the data. The development of a theoretical, model-based procedure capable of accurately predicting psychophysical behavior could alleviate these issues by enabling researchers to rapidly evaluate hypotheses prior to conducting experiments. This approach may also provide additional insight into auditory processing by establishing a link between psychophysical behavior and physiology. Signal detection theory has previously been combined with an auditory model to generate theoretical predictions of psychophysical behavior. Commonly, the ideal processor outperforms human subjects. In order for this model-based technique to enhance the study of auditory processing, discrepancies must be eliminated or explained. In this paper, we investigate the possibility that neural variability, which results from the randomness inherent in auditory nerve fiber responses, may explain some of the previously observed discrepancies. In addition, we study the impact of combining information across nerve fibers and investigate several models of multiple-fiber signal processing. Our findings suggest that neural variability can account for much, but not all, of the discrepancy between theoretical and experimental data.

Acoustic Stimulation↗

Somatosensory, multisensory, and task-related neurons in cortical area 7b (PF) of unanesthetized monkeys.

1. The goal of this study was to quantitatively characterize the response properties of somatosensory and multisensory neurons in cortical area 7b (or PF) of monkeys that were behaviorally trained to perform an appetitive tolerance-escape task. Particular emphasis was given to characterizing nociceptive thermal responses and correlating such responses to thermal pain tolerance as measured by escape frequency. 2. A total of 244 neurons that responded to somatosensory stimulation alone or to both somatosensory and visual stimulation (multisensory) were isolated and studied in the trigeminal region of cortical area 7b. Thirty neurons responded only to visual stimulation. Thermoreceptive neurons formed approximately 13% (31 of 244) of the neurons that had somatosensory response properties. Thermal nociceptive neurons made up approximately 9% (21 of 244) of the neurons that had somatosensory response properties or approximately 68% (21 of 31) of the neurons that had thermoreceptive response properties. Thermal nociceptive neurons responded either exclusively to noxious thermal stimuli (high-threshold thermoreceptive, HTT) or differentially to nonnoxious and noxious thermal stimuli (wide-range thermoreceptive, WRT). Multimodal HTT neurons had nonnociceptive (low-threshold mechanoreceptive, LTM) and/or nociceptive (nociceptive-specific, wide-dynamic-range) mechanical receptive fields, whereas multimodal WRT neurons had only nonnociceptive (LTM) mechanical receptive fields. Thermal nonnociceptive neurons (low-threshold thermoreceptive, LTT) made up approximately 3% (8 of 244) of the neurons that had somatosensory properties or approximately 26% (8 of 31) of the neurons that were thermoreceptive. The background discharge of two thermoreceptive neurons (6%, 2 of 31) was inhibited by innocuous thermal stimulation. 3. Thermal nociceptive neurons (HTT and WRT) were functionally differentiated by statistical analyses into subpopulations that did encode (HTT-EN, WRT-EN) and did not encode (HTT-NE, WRT-NE) the magnitude of noxious thermal stimulus intensities. The mean slopes and median regression coefficients for the stimulus-response (S-R) functions of HTT-EN and WRT-EN neurons, respectively, were significantly greater than those for the S-R functions of HTT-NE and WRT-NE neurons. In contrast to HTT-NE and WRT-NE neurons, HTT-EN and WRT-EN neurons reliably encoded the magnitude of noxious thermal intensity by grading their mean discharge frequency. 4. The S-R functions of HTT-EN and WRT-EN neurons, unlike those of HTT-NE and WRT-NE neurons, closely approximated stimulus intensity-escape frequency functions.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways↗

Derived acoustically evoked brainstem responses by means of narrow-band and notched-noise masking in normal-hearing subjects.

Derived acoustically evoked brainstem responses determined by narrow-band and notched-noise masking at 2000 Hz frequencies and below were investigated and compared with those obtained in the use of the common high-pass masking technique. With all three masking methods a dominant late wave Wa could be detected at less than or equal to 1 600 Hz centre frequencies of the derived band and this wave is typical of the excitation of the apical section of the cochlea. The latency of this wave increases when the centre frequency is decreased at a constant stimulation level. For a constant centre frequency of the derived band the latency of wave Wa increases when the stimulation level of the click and in parallel to that the masking level is decreased. Wave Wa could be found near the hearing threshold in all three masking methods. The notched-noise masking method offers major advantages provided that full masking of the frequency regions above and below the notch frequency range is carried out. A direct frequency-specific stimulation takes place and there is no need for subtracting one response from a second one. This reduces the investigation time and the memory capacity of the measuring device. The advantages and applicability of the notched-noise masking method have to be re-checked in patients with different frequency-dependent loss of hearing.

Adolescent↗

Principles of cross-modal competition: evidence from deficits of attention.

How does the attentional system coordinate the processing of stimuli presented simultaneously to different sensory modalities? We investigated this question with individuals with neurological damage who suffered from deficits of attention. In these individuals, we examined how the processing of tactile stimuli is affected by the simultaneous presentation of visual or auditory stimuli. The investigation demonstrated that two stimuli from different modalities are in competition when attention is directed to the perceptual attributes of both, but not when attention is directed to the perceptual attributes of one and the semantic attributes of the other. These findings reveal a differentiated attentional system in which competition is modulated by the level of stimulus representation to which attention is directed.

Aged↗

Vestibulo-ocular response of human subjects seated in a pivoting support system during 3 Gz centrifuge stimulation.

The vestibulo-ocular reflex (VOR) and spatial orientation perceptions were recorded in 15 subjects during 3 Gz centrifuge runs. These data were obtained to study two issues: (1) to gain insight into reports of asymmetrical disorientation and disturbance during acceleration and deceleration of centrifuge runs like those used to train pilots on the procedures to counteract G-induced loss of consciousness (G-LOC); (2) to study the effects of sustained vertical linear acceleration on the vestibular system. The centrifuge angular velocity profile consisted of a 19 s angular acceleration to 3 Gz that was sustained for 5 min during a period of constant angular velocity, and a 19 s deceleration to 1 Gz. The runs were repeated three times with the subject facing the motion and three times with the subject's back to the motion. The VOR and spatial orientation perceptions from the eight subjects who completed all six runs were analyzed. The total VOR response during acceleration and deceleration was composed of interacting angular (AVOR) and linear components (LVOR). Asymmetries in pitch orientation perception between centrifuge acceleration and decleration were not matched by asymmetries in the total VOR slow phase velocity. During the constant velocity high-G phase of the run, sustained up-beating LVOR (Lz nystagmus) was present in 14 of the 15 subjects. Significant individual differences in Lz nystagmus were found, but the maximum Lz response in our 15 subjects was probably of insufficient magnitude to degrade visual scan of cockpit instruments. Mean magnitudes ranged from 0 to 10 deg/s at 90 s from onset of centrifuge run.

Adult↗

Rhesus macaques spontaneously perceive formants in conspecific vocalizations.

We provide a direct demonstration that nonhuman primates spontaneously perceive changes in formant frequencies in their own species-typical vocalizations, without training or reinforcement. Formants are vocal tract resonances leading to distinctive spectral prominences in the vocal signal, and provide the acoustic determinant of many key phonetic distinctions in human languages. We developed algorithms for manipulating formants in rhesus macaque calls. Using the resulting computer-manipulated calls in a habituation/dishabituation paradigm, with blind video scoring, we show that rhesus macaques spontaneously respond to a change in formant frequencies within the normal macaque vocal range. Lack of dishabituation to a "synthetic replica" signal demonstrates that dishabituation was not due to an artificial quality of synthetic calls, but to the formant shift itself. These results indicate that formant perception, a significant component of human voice and speech perception, is a perceptual ability shared with other primates.

Animals↗

Auditory stimulation alters the pattern of 2-deoxyglucose uptake in the inner ear.

The 2-deoxy-D-glucose (2-DG) autoradiographic technique was adapted for application to the inner ear. The uptake of [14C]-DG during silence was compared with that observed during exposure to wide band noise (WBN) or pure tones at an intensity level of 85 db SPL. In silence, the highest levels of 2-DG uptake were observed in the spiral ligament, spiral prominence and stria vascularis, with approximately equal levels of uptake in each structure. The high levels of 2-DG uptake observed in the ligament and prominence are surprising, and suggest a more active role for these structures in cochlear function than has previously been suspected. Levels of uptake in the organ of Corti, spiral ganglion and VIIIth nerve were much lower, although well above background. During exposure to WBN, 2-DG uptake increased markedly in the VIIIth nerve, and spiral ganglion throughout the cochlea, and in the organ of Corti in the lower basal turn. 2-DG uptake did not change significantly in the spiral ligament or stria vascularis. During pure tone exposure, increased 2-DG uptake was noted in localized regions of the VIIIth nerve and spiral ganglion.

Animals↗