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Neural mechanisms underlying melodic perception and memory for pitch.

The neural correlates of music perception were studied by measuring cerebral blood flow (CBF) changes with positron emission tomography (PET). Twelve volunteers were scanned using the bolus water method under four separate conditions: (1) listening to a sequence of noise bursts, (2) listening to unfamiliar tonal melodies, (3) comparing the pitch of the first two notes of the same set of melodies, and (4) comparing the pitch of the first and last notes of the melodies. The latter two conditions were designed to investigate short-term pitch retention under low or high memory load, respectively. Subtraction of the obtained PET images, superimposed on matched MRI scans, provides anatomical localization of CBF changes associated with specific cognitive functions. Listening to melodies, relative to acoustically matched noise sequences, resulted in CBF increases in the right superior temporal and right occipital cortices. Pitch judgments of the first two notes of each melody, relative to passive listening to the same stimuli, resulted in right frontal-lobe activation. Analysis of the high memory load condition relative to passive listening revealed the participation of a number of cortical and subcortical regions, notably in the right frontal and right temporal lobes, as well as in parietal and insular cortex. Both pitch judgment conditions also revealed CBF decreases within the left primary auditory cortex. We conclude that specialized neural systems in the right superior temporal cortex participate in perceptual analysis of melodies; pitch comparisons are effected via a neural network that includes right prefrontal cortex, but active retention of pitch involves the interaction of right temporal and frontal cortices.

Auditory Perception↗

Pitch, periodicity, and auditory organization.

The perception of pitch forms the basis of musical melody and harmony. It is also among the most precise of all our human senses, and with imagination, this precision can be used experimentally to investigate the functioning of the auditory system. This tutorial presents auditory demonstrations from the zoo of pitch effects: pitch shifts, noise pitch, virtual pitch, dichotic pitch, and the pitches of things that are not there at all. It introduces models of auditory processing, derived from contemporary psychoacoustics and auditory physiology, and tests these models against the experimental effects. It concludes by describing the critical role played by pitch in the important human ability to disentangle overlapping sources of sound.

Cochlea↗

Cochlear implant-mediated perception of music.

PURPOSE OF REVIEW: This paper examines and consolidates recent advances in cochlear implant sound processing from the perspective of music perception, which is increasingly viewed as one of the most difficult of all listening conditions. RECENT FINDINGS: Music is an essentially abstract, complex form of sound composed of multiple layers of sounds that vary in temporal presentation, frequency distribution, and harmonic content. As a result, music perception is perhaps the most challenging aspect of implant-mediated listening. Thus far, implant performance has shown poor performance overall during perception of musical pitches, melodies, and timbre while perception of rhythm is relatively good. Recent advances in implant sound processing strategies, particularly the use of current distribution along adjacent electrodes, have promising early results in terms of improving the number of pitch percepts available to cochlear implant listeners. SUMMARY: Music perception poses auditory challenges that can exceed those of language perception during cochlear implant-mediated listening. These challenges should be emphasized to patients prior to implantation. Although rhythm perception via cochlear implants is reasonably good using simple test paradigms, significant work remains to improve critically important aspects of music perception, including melody and timbre. New implant processing strategies are encouraging and should lead to improved music perception in the near future.

Auditory Perception↗

Automatic discrimination of phonetically relevant and irrelevant vowel parameters as reflected by mismatch negativity.

An auditory event-related brain potential called mismatch negativity (MMN) was measured to study the perception of vowel pitch and formant frequency. In the MMN paradigm, deviant vowels differed from the standards either in F0 or F2 with equal relative steps. Pure tones of corresponding frequencies were used as control stimuli. The results indicate that the changes in F0 or F2 of vowels significantly affected the MMN amplitudes. The only variable significantly affecting the MMN latencies was sex which, however, did not have any effect on the amplitudes of the MMN. As expected, the MMN amplitudes increased with an increase in the acoustical difference between the standards and the deviants in all cases. On the average, the amplitudes were lower for the vowels than for the pure tones of equal loudness. However, in vowels, minor frequency changes in F0 produced higher MMN amplitudes than similar relative changes in F2. It was also noted that even the smallest and phonetically irrelevant change in F2 was detected by the MMN process. In overall, the results demonstrate that the MMN can be measured separately for F0 and F2 of vowels, although the MMN responses show large interindividual differences.

Adult↗

The perception of frequency peaks and troughs in wide frequency modulations.

This work was concerned with the perception of "instantaneous pitch" in continuously frequency modulated sounds. In experiment 1, a 70-dB sinusoidal carrier, close to 1 kHz, was modulated by the exponential of periodic functions corresponding to the sum of a few sinusoids [e.g., sin(at)+sin(3at)]. Each modulation had a fundamental frequency (a/2 pi) of 1.5 Hz and was symmetric on the dimensions of time and log frequency. Thirty listeners identified discrete melodic motifs within these stimuli. The pitches of the identified notes mainly corresponded to the local frequency maxima; generally, the local minima were not heard as auditory "events" (pitch singularities). A similar perceptual asymmetry was not observed for comparable sequences of discrete tones. In experiments 2-4, frequency difference limens were measured for the maxima and minima of continuous frequency modulations, using an adaptive forced-choice method. Sinusoidal carriers were modulated by the exponential of one cycle of a 5-Hz cosine function, starting at phase pi or phase 0 and giving an overall frequency swing of about 0.5 oct. For maxima and minima around 1 kHz, frequency shifts of maxima were better detected than frequency shifts of minima, by an average factor of 2. Generally, this asymmetry did not decrease as a function of subjects' training in the discrimination task, and was still present when frequency minima were given a 6-dB intensity advantage over frequency maxima. No explanation was found for the advantage of frequency maxima with respect to perceptual salience (experiment 1) or discriminability (experiments 2-4).

Adolescent↗

Pitch and timing abilities in adult left-hemisphere-dysphasic and right-hemisphere-damaged subjects.

The production and perception of pitch and rhythm were tested in patients with acquired unilateral left-hemisphere (LH) lesions (and subsequent motor dysphasia, n = 13), patients with unilateral right-hemisphere (RH) lesions (n = 14), and normal age-matched controls. While the LH dysphasic subjects were not generally impaired on the production or perception of pitch, they were grossly impaired on the production and perception of rhythm. The RH subjects, in contrast, were impaired on measures of pitch perception and production, including the discrimination and production of single notes and of melodies. It is concluded that the two hemispheres differ in their specialization for the perception and production of pitch and rhythm.

Adult↗

Production and perception of word tones (pitch accents) in patients with left and right hemisphere damage.

The present paper addresses the question of the functional lateralization of tones in tone languages. Tonal perception and production of right-hemisphere-damaged (RHD) and left-hemisphere-damaged (LHD) speakers of East Norwegian were investigated. East Norwegian is a tone language with an opposition between two tones (pitch accents). The ability to distinguish auditorily between the two accents was normal in the RHD group but reduced in the LHD group. Tonal production was near normal in the RHD group, whereas the LHD group tended to have a production deficit.

Adult↗

Possible influence of linguistic musical background on perceptual pitch-matching tasks: a pilot study.

Linguistic background has been identified as important in the perception of pitch, particularly between tonal versus nontonal languages. In addition, a link between native language and the perception of musical pitch has also been established. This pilot study examined the perception of pitch between listeners from tonal and nontonal linguistic cultures where two different styles of music originate. Listeners were 10 individuals born in China who ranged in age from 25 to 37 years and had spent on the average 30 mo. in the USA and 10 individuals, born on the Indian subcontinent, who ranged in age from 22 to 31 years, and had spent an average of 13 mo. in the USA. Listeners from both groups participated in two conditions. One condition involved listening to a selection of music characteristic of the individual's culture (China, pentatonic scale; Indian subcontinent, microtones), and one condition involved no music. All listeners within each condition participated in two voice pitch-matching tasks. One task involved matching the lowest and highest pitch of tape-recorded voices to a note on an electronic keyboard. Another task involved matching the voice pitch of tape-recorded orally read words to a note on the keyboard. There were no differences between the two linguistic groups. Methodological limitations preclude generalization but provide the basis for further research.

Adolescent↗

Frequency and frequency-ratio resolution by possessors of absolute and relative pitch: examples of categorical perception.

The methodology derived from the trace-context theory of intensity resolution (Durlach and Braida, 1969; Macmillan et al., 1988) was applied to resolution over an octave range along two continua: a sequential-frequency-ratio continuum for possessors of relative pitch (RP), and a pure-tone frequency continuum for possessors of absolute pitch (AP). The performance of both RP and AP possessors was exceptional in that total identification sensitivity along both continua was much greater than identification sensitivity along unidimensional psychophysical continua characterized by the 7 +/- 2 rule. In addition, the performance of RP possessors was exceptional in that, on average, total sensitivity for identification resolution was greater than sensitivity for resolution in discrimination. Finally, identification sensitivity between category prototypes (chromatic semits) along both continua was approximately the same as identification sensitivity between phonemic category prototypes along speech continua, despite the fact that both the discrimination ranges and the total number of categories are much larger for the two pitch continua.

Humans↗

Adaptation of residue pitch.

Residue pitch was examined by means of adaptation. The monaural perceptibility of a residue pitch was measured before and after exposure to ipsilateral or contralateral adapatation stimuli of variable spectral composition. Several aspects of adaptation were investigated: pitch channel specificity, number of stimulus components necessary to adapt a residue channel, importance of temporal and spectral factors for adaptation, and the relation between pure-tone and residue pitch. It was found that there are residue-pitch channels specifically sensitive to the spectral region and ear of presentation of the stimulus. At least two harmonics were necessary to adapt a residue channel, and pitch channels were found to be insensitive to temporal factors. Residue and pure-tone pitch seemed to be extracted by different mechanisms, having independent channels. The data presented augment the understanding of pitch perception and suggest refinements for pitch-perception models.

Cues↗

Perceived pitch of whispered vowels--relationship with formant frequencies: a preliminary study.

To clarify the role of formant frequency in the perception of pitch in whispering, we conducted a preliminary experiment to determine (1.) whether speakers change their pitch during whispering; (2.) whether listeners can perceive differences in pitch; and (3.) what the acoustical features are when speakers change their pitch. The listening test of whispered Japanese speech demonstrates that one can determine the perceived pitch of vowel /a/ as ordinary, high, or low. Acoustical analysis revealed that the perception of pitch corresponds to some formant frequencies. Further data with synthesized whispered voice are necessary to confirm the importance of the formant frequencies in detail for perceived pitch of whispered vowels.

Adult↗

A comparative study on the effect of pure-tone exposure of the guinea pig cochlea.

Electrophysiological methods were applied to 160 healthy adult male guinea pigs in order to investigate the effects of pure-tone exposure for 24 h on the inner ear. A reduction in cochlear microphonics (CM), action potential (AP) and endocochlear potential was observed following exposure to 110 dB at 100 Hz, 100 dB at 200 and 600 Hz and 95 dB at 2 kHz. The observed K+ endolymphatic concentration during 40 min anoxia remained unchanged. In contrast K+ decreased in control animals and following exposure to pure tones varying from 110 dB at 60 Hz to 85 dB at 2 kHz. These findings indicate that high frequency tones have a greater effect on inner ear functions than those of lower frequency, decreasing the maximum output voltage of CM and AP but not changing K+ endolymphatic concentration.

Acoustic Stimulation↗

Detection of frequency modulation in the FM-bat Phyllostomus discolor.

In a two-alternative forced-choice procedure lesser spear-nosed bats, Phyllostomus discolor, had to discriminate between a pure tone stimulus and a sinusoidally frequency-modulated signal generated at the same carrier frequency as the tone. Modulation depths of the SFM stimuli were reduced until the animals' performance dropped below the 75%-correct level which was used to determine difference limens for detection of frequency modulation (FMDL). The dependence of FMDLs on modulation and carrier frequency was systematically investigated. For a carrier frequency of 18.5 kHz, average FMDLs increased from 95 Hz at a modulation frequency of 10 Hz to 820 Hz at a modulation frequency of 2000 Hz which corresponds to Weber ratios (2 delta f/f) of 0.005 and 0.044 respectively. Further, difference limens were found to increase linearly in proportion to carrier frequency throughout a major part (9-74 kHz) of the species' hearing range. In comparison to other mammals, P. discolor has a pronounced capability for frequency discrimination which might be related to the extensive use of individually distinct frequency-modulated communication calls and audio-vocal learning.

Acoustic Stimulation↗

Thalamocortical transformation of responses to complex auditory stimuli.

In unanesthetized guinea pigs, thalamic (CGM), and cortical (auditory I) neurons were recorded simultaneously. Nine of 69 neuron pairs showed a positive cross-correlation of their spontaneous activities, with increased discharge probability of the cortical neuron beginning 2--5 ms after the discharge of the CGM-neuron. The individual neurons of such pairs had an identical CF and the same spectral responsiveness. The responses of cortical neurons to pure tones were much more phasic than those of the corresponding CGM-neurons. Thalamic neurons could be driven up to much higher AM- and FM-modulation frequencies (100 Hz) than cortical neurons, which usually ceased to follow AM-frequencies above 20 Hz. Stronger or weaker suppression of tonic response components in cortical and thalamic neurons and the lower AM-range of cortical neurons is related to stronger or weaker intracortical and intrathalamic inhibition respectively. Response characteristics to FM-stimuli are similar to those of AM-stimuli. All CGM and cortical neurons responded to a variety of natural calls of the same or of other species. Responses of CGM-cells represented more components of a call than cortical cells even if the two cells were synaptically connected. In cortical cells, repetitive elements of a call were not represented if the repetition rate was too high. High modulation frequencies within a call, such as those of the fundamental frequency, could still be separated in the response of some CGM-neurons, but never in those of cortical neurons. Both CGM and cortical cells responded essentially to transients (amplitude or frequency modulations) within a call, if spectral components of such elements were within the spectral sensitivity of the cell. Spectral components outside the spectral sensitivity range could result in suppression of spontaneous discharge rate. Responses of cortical and CGM-cells, and thus the representation of call elements by neuronal responses, varied with the intensity of a call. It is suggested that, at higher levels of the auditory system, essential information about the temporal features of complex sounds may be represented by neural responses to transients in various spectral regions.

Animals↗