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Musical and linguistic processing in song perception.

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.

Acoustic Stimulation↗

Dynamics of precise spike timing in primary auditory cortex.

Although single units in primary auditory cortex (A1) exhibit accurate timing in their phasic response to the onset of sound (precision of a few milliseconds), paradoxically, they are unable to sustain synchronized responses to repeated stimuli at rates much beyond 20 Hz. To explore the relationship between these two aspects of cortical response, we designed a broadband stimulus with a slowly modulated spectrotemporal envelope riding on top of a rapidly modulated waveform (or fine structure). Using this stimulus, we quantified the ability of cortical cells to encode independently and simultaneously the stimulus envelope and fine structure. Specifically, by reverse-correlating unit responses with these two stimulus dimensions, we measured the spectrotemporal response fields (STRFs) associated with the processing of the envelope, the fine structure, and the complete stimulus. A1 cells respond well to the slow spectrotemporal envelopes and produce a wide variety of STRFs. In over 70% of cases, A1 units also track the fine-structure modulations precisely, throughout the stimulus, and for frequencies up to several hundred Hertz. Such a dual response, however, is contingent on the cell being driven by both fast and slow modulations, in that the response to the slowly modulated envelope gates the expression of the fine structure. We also demonstrate that either a simplified model of synaptic depression and facilitation, and/or a cortical network of thalamic excitation and cortical inhibition can account for major trends in the observed findings. Finally, we discuss the potential functional significance and perceptual relevance of these coexistent, complementary dynamic response modes.

Acoustic Stimulation↗

Micromechanics of the reptilian ear.

This paper reviews selected physiology and anatomy of the lizard peripheral auditory system and presents the implications of these data for understanding cochlear micromechanics. The reasons for studying the micromechanics of the lizard ear are presented along with the difficulties in conducting such studies in the mammal. Data from the Anguid and Iguanid families of lizards demonstrate the influence of the hair cell system on neural tuning. In addition, evidence from the Iguanid suggests that cilium length may help determine neural tonotopic organization. In general, the reptilian research presented shows the importance of the hair cell system for determining the tuning of associated nerve fibers and in understanding the transduction process in general.

Animals↗

Neurone response latency in the inferior colliculus in relation to the auditory brainstem responses (ABR) in the guinea pig.

In order to determine the possible source of the later vertex positive waves of the guinea pig auditory brainstem response (ABR), we have measured field potentials on the surface and at depths through inferior colliculus whilst making concurrent recordings of single unit responses (using identical broadband click stimuli). We find neurones, restricted to the antero-ventral inferior colliculus, with characteristic frequencies in excess of 12 kHz which have well synchronized onset responses short enough (c. 5 ms) to contribute to the P5 wave of the ABR.

Animals↗

Cortical processing of musical consonance: an evoked potential study.

Cortical processes underlying perception of musical consonance were investigated by long-latency auditory evoked potentials (EPs). Subjects listened to a random sequence of dyadic pure tones paired at various pitch intervals (1, 4, 6, 7, or 9 semitones). Amplitudes of P2 and N2 components of auditory EPs were significantly modulated by pitch interval of the dyads, being most negative for 1 semitone (minor second) and least negative or most positive for 7 semitones (perfect fifth). The results indicate that neural processing of consonance depend not only on peripheral mechanisms in the inner ear but also on higher associative processing of pitch relationships in the cerebral cortex.

Acoustic Stimulation↗

Multiresolution spectrotemporal analysis of complex sounds.

A computational model of auditory analysis is described that is inspired by psychoacoustical and neurophysiological findings in early and central stages of the auditory system. The model provides a unified multiresolution representation of the spectral and temporal features likely critical in the perception of sound. Simplified, more specifically tailored versions of this model have already been validated by successful application in the assessment of speech intelligibility [Elhilali et al., Speech Commun. 41(2-3), 331-348 (2003); Chi et al., J. Acoust. Soc. Am. 106, 2719-2732 (1999)] and in explaining the perception of monaural phase sensitivity [R. Carlyon and S. Shamma, J. Acoust. Soc. Am. 114, 333-348 (2003)]. Here we provide a more complete mathematical formulation of the model, illustrating how complex signals are transformed through various stages of the model, and relating it to comparable existing models of auditory processing. Furthermore, we outline several reconstruction algorithms to resynthesize the sound from the model output so as to evaluate the fidelity of the representation and contribution of different features and cues to the sound percept.

Acoustic Stimulation↗

Three-dimensional baselines for perceived self-motion during acceleration and deceleration in a centrifuge.

Three-dimensional motion trajectories were computed, representing the motions that would be perceived by a perfect processor of acceleration information during the acceleration and deceleration stages of a centrifuge run. These motions serve as "baselines" for perceived self-motion in a centrifuge, and depend on the initial perception of orientation and velocity immediately preceding the acceleration and immediately preceding the deceleration. The baselines show that a perfect processor of acceleration information perceives self-motion during centrifuge deceleration significantly differently from self-motion during centrifuge acceleration, despite the fact that the angular accelerations have equal magnitude (with opposite direction). At the same time, the baselines can be compared with subjects' reported perceptions to highlight limitations of the nervous system; limitations and peculiarities of the nervous system are identified as deviations from a baseline. As a result, peculiarities of the nervous system are held responsible for any perception of pitch or roll angular velocity or change in tilt of the body-horizontal plane of motion during the centrifuge run. On the other hand, baselines explain perception of tilt position during deceleration, linear velocity, possible lack of significant linear velocity during deceleration, and yaw angular velocity, including on-axis angular velocity during centrifuge deceleration. The results lead to several experimental questions.

Acceleration↗

Development of inferior colliculus response properties in C57BL/6J mouse pups.

Response properties of inferior colliculus (IC) neurons were studied in traquilized C57BL/6J mice during a period of rapid auditory system development between 12 and 17 days of age. In IC units of the youngest mice, spontaneous activity was absent, a disproportionate number of onset responses was observed, and many units were not securely driven by sound. Frequency response ranges were restricted to relatively low frequencies, sharpness of tuning was poor, and thresholds at best frequencies (BFs) were quite high. Dynamic intensity ranges were restricted, but nonmonotonic functions were observed. By 15--17 days of age, spontaneous activity was appreciable, incidences of response patterns were near adult proportions, and most units in the ventrolateral nucleus were securely driven by tones. Response ranges had expanded markedly to include high frequencies, sharpness of tuning increased, and thresholds had decreased. Dynamic intensity ranges and intensity functions were similar to those observed in adult mice.

Acoustic Stimulation↗

Similarity as an organising principle in short-term memory.

The role of stimulus similarity as an organising principle in short-term memory was explored in a series of seven experiments. Each experiment involved the presentation of a short sequence of items that were drawn from two distinct physical classes and arranged such that item class changed after every second item. Following presentation, one item was re-presented as a probe for the 'target' item that had directly followed it in the sequence. Memory for the sequence was considered organised by class if probability of recall was higher when the probe and target were from the same class than when they were from different classes. Such organisation was found when one class was auditory and the other was visual (spoken vs. written words, and sounds vs. pictures). It was also found when both classes were auditory (words spoken in a male voice vs. words spoken in a female voice) and when both classes were visual (digits shown in one location vs. digits shown in another). It is concluded that short-term memory can be organised on the basis of sensory modality and on the basis of certain features within both the auditory and visual modalities.

Auditory Perception↗

Implicit investigations of tonal knowledge in nonmusician listeners.

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.

Acoustic Stimulation↗

Some current concepts of cochlear mechanics.

The development of the current concepts is reviewed in historical perspective. Helmholtz's hypothesis of basilar-membrane resonance was partially confirmed and partially defeated by Békésy's experiments on models and postmortem cochlear preparations. He discovered that sound was propagated in the cochlea in the form of traveling waves which reached a flat maximum at a frequency-dependent location. Mathematical theory explained this type of sound propagation as a special case of surface waves. Johnstone and his coworkers discovered that the maximum of cochlear vibration in living animals was much sharper than postmortem, and more recently Khanna and Johnstone independently determined the maximum to be nearly as sharp as the tuning curves of the inner hair cells and the auditory-nerve fibers. These findings, together with the work at the Massachusetts Institute of Technology on alligator lizards, have led to new concepts of cochlear mechanics which include hypothesized micromechanical processes in the organ of Corti. These concepts deal not only with the sharpness of basilar-membrane tuning but also with the details of the basilar-membrane amplitude and phase characteristics, as well as with the hair cell and neural tuning curves and response phases. They suggest that some sharpening of the tuning curves occurs between the basilar membrane and hair-cell responses. Such sharpening has been demonstrated in lizards, but in the mammalian ear, the relation is less clear.

Animals↗

A simple model of cochlear micromechanics in the mammal and lizard.

A model is presented which is a simple representation of cochlear micromechanics in the lizard and mammal. The model is a linear, time-invariant mechanical system, consisting of two coupled mechanical filters. It is capable of a sharply tuned first and second filter, consistent with the sharply tuned basilar membrane and auditory nerve fibers in the mammal. When parameters are adjusted for the alligator lizard (Gerrhonotus multicarinatus), the model generates a sharply tuned second filter without a sharply tuned first filter. In the alligator lizard, the auditory nerve fibers are sharply tuned but the basilar membrane is not. The model supports the hypothesis that cochlear transduction is not fundamentally different in the mammal and the alligator lizard.

Animals↗

On cross-modal similarity: the perceptual structure of pitch, loudness, and brightness.

Examined how pitch and loudness correspond to brightness. In the Experiment 1, 16 Ss identified which of 2 lights more resembled each of 16 tones; in Experiment 2, 8 of the same 16 Ss rated the similarity of lights to lights, tones to tones, and lights to tones. (1) Pitch and loudness both contributed to cross-modal similarity, but for most Ss pitch contributed more. (2) Individuals differed as to whether pitch or loudness contributed more; these differences were consistent across matching and similarity scaling. (3) Cross-modal similarity depended largely on relative stimulus values. (4) Multidimensional scaling revealed 2 perceptual dimensions, loudness and pitch, with brightness common to both. A simple quantitative model can describe the cross-modal comparisons, compatible with the view that perceptual similarity may be characterized through a malleable spatial representation that is multimodal as well as multidimensional.

Attention↗

Lateralization of phonetic and pitch discrimination in speech processing.

Cerebral activation was measured with positron emission tomography in ten human volunteers. The primary auditory cortex showed increased activity in response to noise bursts, whereas acoustically matched speech syllables activated secondary auditory cortices bilaterally. Instructions to make judgments about different attributes of the same speech signal resulted in activation of specific lateralized neural systems. Discrimination of phonetic structure led to increased activity in part of Broca's area of the left hemisphere, suggesting a role for articulatory recoding in phonetic perception. Processing changes in pitch produced activation of the right prefrontal cortex, consistent with the importance of right-hemisphere mechanisms in pitch perception.

Acoustic Stimulation↗

Response properties of auditory activated cells in the occipital cortex of the blind mole rat: an electrophysiological study.

Previous studies have demonstrated that despite its blindness, the subterranean blind mole rat (Spalax ehrenbergi) possesses a noticeable lateral geniculate nucleus and a typical cyto-architectural occipital cortex that are reciprocally connected. These two areas, as revealed by the metabolic tracer 2-deoxyglucose, are activated by auditory stimuli. Using single unit recordings, we show that about 57% of 325 cells located within the occipital cortex of anesthetized mole rats responded to at least one of the following auditory stimuli--white noise, pure tones, clicks, and amplitude modulated tones--with the latter two being the most effective. About 85% of cells driven by either contralateral or ipsilateral stimulation also responded to binaural stimulation; about 13% responded only to binaural stimulation; and 2% were driven exclusively by contralateral stimulation. Comparing responsiveness and response strength to these three modes of stimulation revealed a contralateral predominance. Mean latency (+/-SD) of ipsilateral and contralateral responses were 48.5+/-32.6 ms and 33.5+/-9.4 ms, respectively. Characteristic frequencies could be divided into two distinct subgroups ranging between 80 and 125 Hz and between 2,500 and 4,400 Hz, corresponding to the most intensive spectral components of the vibratory intraspecific communication signals and airborne vocalizations.

Acoustic Stimulation↗

Brain potentials in patients with music perception deficits: evidence for an early locus.

Twelve patients with an acute cerebrovascular accident were assigned to a group with music perception deficits (amusia, n = 6) or a group without such deficits (n = 6) on the basis of a new test-battery for music-perception skills. Event-related brain potentials (ERPs) were recorded in an auditory classification task designed to elicit several components; the N1 as a correlate of initial auditory cortical processing, the P3a as an index of automatic attentional orienting, and the P3b as a measure for controlled stimulus evaluation. Patients with amusia showed a significant amplitude decrement for the P3a relative to controls and patients without amusia suggesting an impairment of early stimulus evaluation. P3b was reduced in both patient groups relative to control. These data show that amusia is quite common in unselected stroke patients and suggest deficits of generic rather than music-specific cognitive processes as the underlying cause.

Acoustic Stimulation↗

Cognitive basis of hallucinations in schizophrenia: role of top-down information processing.

Hallucinations in schizophrenia have been regarded to result from the erroneous attribution of internally generated information to an external source. Distortions in mental imagery may underlie such confusions. We investigated performance of 77 subjects on multiple behavioral measures of auditory and visual mental imagery and perception, and a measure of reality monitoring. Comparisons were made between performance of schizophrenia patients with (N=22) and without (N=35) hallucinations and matched normal comparison subjects (N=20), after controlling for attentional factors. No differences emerged on any of the mental imagery measures, nor on reality monitoring accuracy. This suggests that there is no stable disposition towards abnormal mental imagery associated with hallucinations. However, for patients with active hallucinations (N=12), hallucination severity correlated positively with a measure of imagery-perception interaction in the auditory modality, r=0.70, p=0.01. Although preliminary, this finding is consistent with recent theoretical proposals in which hallucinations have been suggested to result from an increased influence of top-down sensory expectations on conscious perception.

Adult↗

Stages of auditory feature conjunction: an event-related brain potential study.

Auditory event-related brain potentials (ERPs) were recorded to tones of different frequencies and locations in a dichotic selective attention task in which Ss responded to occasional deviant tones of a prespecified location and frequency. Attention effects were isolated as negative difference (Nd) waves by subtracting ERPs to tones with no attended features from ERPs to the same tones when they shared target frequency, location, or both cues. The N1/P90 (latency 80-100 ms), originating in a tonotopically organized generator, was enhanced for all tones in the attended ear. Nd waves, beginning at 80 ms and lasting up to 700 ms, were seen to tones with either attended feature. Nd waves to frequency and location features had different scalp distributions consistent with generation in different cortical fields. Conjunction-specific Nds began 30-50 ms after Nds to individual features. The relative timing suggests that feature conjunction began before the analysis of individual features was complete.

Acoustic Stimulation↗