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Response- and stimulus-related ERP asymmetries in a tonal oddball task: a Laplacian analysis.

Previous studies have found greater P3 amplitude over right than left hemisphere sites in a tonal oddball task with a reaction time (RT) response. This asymmetry had a central topography, and interacted with response hand. Identification of the processes underlying these asymmetries requires the use of additional methods for separating response- and stimulus-related contributions. We applied local Hjorth and spherical spline algorithms to compute surface Laplacian topographies of ERP data recorded from 30 scalp electrodes in a pooled sample of 46 right-handed healthy adults. For both methods, the current sources underlying the late positive complex were largest at medial parietal regions, but were asymmetric at central and frontocentral sites. Although a frontocentral sink contralateral to the response hand contributed to the asymmetry of the classic P3 peak, the source asymmetry was most robust after the sink had resolved. The late source was largest at electrode C4 for right hand responses, and was further enhanced in subjects showing a dichotic left ear advantage, but was unrelated to response speed. We conclude that the right hemisphere source reflects an interaction of response-related asymmetries with right hemisphere processes responsible for pitch discrimination.

Acoustic Stimulation↗

Perception of structure in short melodic sequences.

Three experiments studied the perception of tone sequences having various degrees of musical structure. Ratings of perceived structure and ease of recognition in transposition were both influenced by harmonic progression (as defined by music theory), the contour (directional changes in pitch), and the excursion or repetition pattern within the sequence. The relation between the original and transposed sequence also affected ease of recognition in accordance with the number of tones shared between the two sequences. The results are described in terms of the abstraction and analysis of levels of pitch relations, an analysis conducted even by musically untrained listeners. The conceptual framework emphasizes the application of musical rules as an illustration of rules governing auditory sequences in general.

Humans↗

Demonstration of presbycusis across repeated measures in a nonhuman primate species.

The progress of age-related hearing loss in three 31-yr-old, two 24-yr-old, and two 9-yr-old rhesus monkeys (Macaca mulatta) was repeatedly examined over a period of 3 yr. Pure-tone audiograms for seven frequencies, .125, .500, 2.000, 4.000, 16.000, 22.667, and 32.000 kHz, were obtained through the use of the psychophysical tracking method. Analysis indicated the presence of presbycusis in the older animals. Animals demonstrating presbycusis showed a progressive decrement in the ability to detect higher frequencies and an overall hearing loss at all frequencies.

Aging↗

Attentional modulation of human auditory cortex.

Attention powerfully influences auditory perception, but little is understood about the mechanisms whereby attention sharpens responses to unattended sounds. We used high-resolution surface mapping techniques (using functional magnetic resonance imaging, fMRI) to examine activity in human auditory cortex during an intermodal selective attention task. Stimulus-dependent activations (SDAs), evoked by unattended sounds during demanding visual tasks, were maximal over mesial auditory cortex. They were tuned to sound frequency and location, and showed rapid adaptation to repeated sounds. Attention-related modulations (ARMs) were isolated as response enhancements that occurred when subjects performed pitch-discrimination tasks. In contrast to SDAs, ARMs were localized to lateral auditory cortex, showed broad frequency and location tuning, and increased in amplitude with sound repetition. The results suggest a functional dichotomy of auditory cortical fields: stimulus-determined mesial fields that faithfully transmit acoustic information, and attentionally labile lateral fields that analyze acoustic features of behaviorally relevant sounds.

Acoustic Stimulation↗

Midfrequency dysfunction in listeners having high-frequency sensorineural hearing loss.

Two-tone unmasking, psychophysical tuning curves and pure-tone masking patterns were measured at 500 and 1000 Hz in 17 listeners having high-frequency sensorineural hearing loss due to noise exposure. Results were compared to similar data obtained from 20 normal-hearing young adults. In addition, measures of word-recognition ability were obtained in quiet and in noise for both groups. The primary findings were as follows: (a) 29% (n = 5) of the hearing-impaired subjects exhibited abnormal results on at least one of the psychoacoustic tasks investigated; (b) the observed abnormalities were reliable; and (c) there appeared to be a relation between the presence of midfrequency dysfunction and degree of difficulty on word-recognition tasks. These results and their implications are discussed.

Adult↗

Neural mechanism for binaural pitch perception via ghost stochastic resonance.

We present a physiologically plausible binaural mechanism for the perception of the pitch of complex sounds via ghost stochastic resonance. In this scheme, two neurons are driven by noise and a different periodic signal each (with frequencies f(1)=kf(0) and f(2)=(k+1)f(0), where k>1), and their outputs (plus noise) are applied synaptically to a third neuron. Our numerical results, using the Morris-Lecar neuron model with chemical synapses explicitly considered, show that intermediate noise levels enhance the response of the third neuron at frequencies close to f(0), as in the cases previously described of ghost resonance. For the case of an inharmonic combination of inputs (f(1)=kf(0)+Deltaf and f(2)=(k+1)f(0)+Deltaf) noise is also seen to enhance the rates of most probable spiking for the third neuron at a frequency f(r)=f(0)+[Deltaf(k+12)]. In addition, we show that similar resonances can be observed as a function of the synaptic time constant. The suggested ghost-resonance-based stochastic mechanism can thus arise either at the peripheral level or at a higher level of neural processing in the perception of pitch.

Biophysics↗

"Seeing things in my head": a synesthete's images for music and notes.

We explored synesthetic experiences (photisms) elicited by music and notes for GS, a 22-year-old female college student, who is a genuine digit synesthete (Mills et al 1999 Cognitive Neuropsychology 16 181-191). After extensive interviews, we conducted five studies using synthesizer notes in different octaves and in different instrument sounds. GS described, drew, or selected her photisms from alternatives. The results were: (a) her photisms had shape, color, movement, and texture: (b) different instruments had photisms with different shapes and families of instruments had similar shapes; (c) low-pitched notes were darker and larger than high notes; (d) instrument and pitch affected each other; (e) a given note was perceived as darker after a high note than after a low note; and (f) GS's drawings of notes became more consistent with repeated trials. Theoretical observations are discussed on the basis of these results.

Acoustic Stimulation↗

Correct tonotopic representation is necessary for complex pitch perception.

The ability to extract a pitch from complex harmonic sounds, such as human speech, animal vocalizations, and musical instruments, is a fundamental attribute of hearing. Some theories of pitch rely on the frequency-to-place mapping, or tonotopy, in the inner ear (cochlea), but most current models are based solely on the relative timing of spikes in the auditory nerve. So far, it has proved to be difficult to distinguish between these two possible representations, primarily because temporal and place information usually covary in the cochlea. In this study, "transposed stimuli" were used to dissociate temporal from place information. By presenting the temporal information of low-frequency sinusoids to locations in the cochlea tuned to high frequencies, we found that human subjects displayed poor pitch perception for single tones. More importantly, none of the subjects was able to extract the fundamental frequency from multiple low-frequency harmonics presented to high-frequency regions of the cochlea. The experiments demonstrate that tonotopic representation is crucial to complex pitch perception and provide a new tool in the search for the neural basis of pitch.

Cochlear Implants↗

Physical basis of two-tone interference in hearing.

The cochlea uses active amplification to capture faint sounds. It has been proposed that the amplifier comprises a set of self-tuned critical oscillators: each hair cell contains a force-generating dynamical system that is maintained at the threshold of an oscillatory instability, or Hopf bifurcation. While the active response to a pure tone provides frequency selectivity, exquisite sensitivity, and wide dynamic range, its intrinsic nonlinearity causes tones of different frequency to interfere with one another in the cochlea. Here we determine the response to two tones, which provides a framework for understanding how the ear processes the more complex sounds of speech and music. Our calculations of two-tone suppression and the spectrum of distortion products generated by a critical oscillator accord with experimental observations of basilar membrane motion and the nervous response. We discuss how the response of a set of self-tuned oscillators, covering a range of characteristic frequencies, represents the structure of a complex sound. The frequency components of the stimulus can be inferred from the timing of neural spikes elicited by the vibrating hair cells. Passive prefiltering by the basilar membrane improves pitch discrimination by reducing interference between tones. Our analysis provides a general framework for examining the relation between the physical nature of the peripheral detection apparatus and psychophysical phenomena such as the sensation of dissonance and auditory illusions.

Cochlea↗

Characterization of deficits in pitch perception underlying 'tone deafness'.

Congenital amusia is a disorder characterized by life-long, selective deficits in the perception of music. This study examined pitch-perception abilities in a group of 10 adults with this disorder. Tests were administered that assessed fine-grained pitch perception by determining thresholds both for the detection of continuous and segmented pitch changes, and for the recognition of pitch direction. Tests were also administered that assessed the perception of more complex pitch patterns, using pitch-sequence comparison tasks. In addition, the perceptual organization of pitch was also examined, using stream segregation tasks that assess the assignment of sounds differing in pitch to one or two distinct perceptual sources. In comparison with 10 control subjects, it was found that the participants with congenital amusia exhibited deficits both at the level of detecting fine-grained differences in pitch, and at the level of perceiving patterns in pitch. In contrast, no abnormalities were identified in the perceptual organization of pitch. The pitch deficits identified are able to account for the music perception difficulties in this disorder, and implicate deficient cortical processing.

Acoustic Stimulation↗

Neuromagnetic evidence for a pitch processing center in Heschl's gyrus.

There have been several attempts to use the neuromagnetic response to the onset of a tonal sound (N100m) to study pitch processing in auditory cortex. Unfortunately, a large proportion of the N100m is simply a response to the onset of sound energy, independent of whether the sound produces a pitch. The current study describes a novel stimulus paradigm designed to circumvent the energy-onset response and thereby isolate the response of those neural elements specifically involved in pitch processing. The temporal resolution of magnetoencephalography enables us to show that the latency and amplitude of this pitch-onset response (POR) vary with the pitch and pitch strength of the tone. The spatial resolution is sufficient to show that its source lies somewhat anterior and inferior to that of the N100m, probably in the medial part of Heschl's gyrus.

Acoustic Stimulation↗

Amplitude and frequency-modulated stimuli activate common regions of human auditory cortex.

Hall et al. (Hall et al., 2002, Cerebral Cortex 12:140-149) recently showed that pulsed frequency-modulated tones generate considerably higher activation than their unmodulated counterparts in non-primary auditory regions immediately posterior and lateral to Heschl's gyrus (HG). Here, we use fMRI to explore the type of modulation necessary to evoke such differential activation. Carrier signals were a single tone and a harmonic-complex tone, with a 300 Hz fundamental, that were modulated at a rate of 5 Hz either in frequency, or in amplitude, to create six stimulus conditions (unmodulated, FM, AM). Relative to the silent baseline, the modulated tones, in particular, activated widespread regions of the auditory cortex bilaterally along the supra-temporal plane. When compared with the unmodulated tones, both AM and FM tones generated significantly greater activation in lateral HG and the planum temporale, replicating the previous findings. These activation patterns were largely overlapping, indicating a common sensitivity to both AM and FM. Direct comparisons between AM and FM revealed a higher magnitude of activation in response to the variation in amplitude than in frequency, plus a small part of the posterolateral region in the right hemisphere whose response was specifically AM-, and not FM-, dependent. The dominant pattern of activation was that of co-localized activation by AM and FM, which is consistent with a common neural code for AM and FM within these brain regions.

Acoustic Stimulation↗

The P3a wave of the auditory event-related potential reveals registration of pitch change during sufentanil anesthesia for cardiac surgery.

BACKGROUND: The N1 and P3 waves of the auditory event-related potential provide information on consciousness and cortical function. The N1 wave is reduced during states of low vigilance. The P3 wave occurs only for stimuli that somehow capture the subject's attention. There are two types of P3:P3a and P3b. The P3a predominates frontally and probably occurs when the subject simply notices the stimulus. The P3b predominates parietally and indicates conscious awareness of the evoking stimulus. The N1 and P3 were recorded in 12 patients during cardiac surgery under sufentanil anesthesia to search for unintentional awareness. The study was limited to the period before cardiopulmonary bypass. METHODS: After premedication with diazepam, morphine, and scopolamine, sufentanil was used for induction (mean dose, 7.9 micrograms/kg) and maintenance (4 micrograms/kg) of anesthesia. No other anesthetics were administered. Recordings were obtained before induction, during induction after loss of consciousness, after tracheal intubation before incision, and before cardiopulmonary bypass. RESULTS: The N1 was attenuated significantly by sufentanil but was not abolished. The P3b occurred only during preinduction. There was no P3 during induction. There was a P3a during postintubation and precardiopulmonary bypass. CONCLUSIONS: The attenuation of N1 from induction onward reflects a decrease in the level of arousal caused by sufentanil. A P3a during postintubation and precardiopulmonary bypass indicates that pitch discrimination at the cortical level occurs but does not prove that conscious awareness has occurred. Whether or not the P3a reflects the regaining of consciousness is not known.

Acoustic Stimulation↗

Neural correlates of absolute pitch differ between blind and sighted musicians.

Several reports have indicated a higher incidence of absolute pitch in blind than in sighted musicians. Employing a pitch memory task, we examined whether a blind absolute pitch musician would rely on different neural correlates than a group of sighted absolute pitch musicians. The blind musician showed significantly more activation of bihemispheric visual association areas, lingual gyrus, parietal and frontal areas than the sighted musicians. Sighted musicians showed more activation of the right primary auditory cortex and the cerebellum when compared with the blind musician. These differences in the activation pattern suggest the use of a different neural network including visual association areas while performing pitch categorization and identification in this blind musician in comparison with sighted musicians.

Acoustic Stimulation↗

Perception of timbral analogies.

Recent studies have investigated the structure of perceptual relations among musical instrument timbres by multidimensional scaling (MDS) techniques. These studies have employed both acoustically produced tones and digitally synthesized imitations and hybrids of acoustic instrument tones. The analyses of dissimilarity ratings for all pairs of a set of tones are usually represented as geometrical structures in a two- or three-dimensional Euclidean space in which the shared 'perceptual' axes are shown to have a qualitative correspondence to acoustic properties such as spectral energy distribution, onset characteristics and degree of change in spectral distribution over the duration of the tone. The present study took as a point of departure a MDS analysis for complex, synthetic tones with the aim of testing whether musician and non-musician listeners used the relations defined by the perceptual space to perform an analogies task of the sort: timbre A is to timbre B as timbre C is to which of two possible timbres, D or D'? A parallelogram model was used to select the D timbres: if the relation between A and B is represented as a vector with both magnitude and direction components, then the appropriate D should form a vector with C having similar magnitude and direction in the timbre space. Aside from conceptual difficulties with the task for both non-musicians and composers, choices for both groups provide support for the parallelogram model indicating a capacity in listeners to perceive abstract relations among the timbres of complex sounds without specific training in such a task.

Acoustic Stimulation↗