[P2 evaluation] Articles

Choisir deux articles dans la liste, provenant de deux intervenants différents. N'oubliez pas de me préciser lequel est pour l'oral et lequel est pour l'écrit. Vu que Barbara Tillmann ne pourra pas assister à l'oral, il serait préférable de choisir ses articles pour l'écrit.


ALG1 : Nat Neurosci. 2005 Mar;8(3):389-95.

Hierarchical and asymmetric temporal sensitivity in human auditory cortices.

Boemio A, Fromm S, Braun A, Poeppel D.

Lateralization of function in auditory cortex has remained a persistent puzzle. Previous studies using signals with differing spectrotemporal characteristics support a model in which the left hemisphere is more sensitive to temporal and the right more sensitive to spectral stimulus attributes. Here we use single-trial sparse-acquisition fMRI and a stimulus with parametrically varying segmental structure affecting primarily temporal properties. We show that both left and right auditory cortices are remarkably sensitive to temporal structure. Crucially, beyond bilateral sensitivity to timing information, we uncover two functionally significant interactions. First, local spectrotemporal signal structure is differentially processed in the superior temporal gyrus. Second, lateralized responses emerge in the higher-order superior temporal sulcus, where more slowly modulated signals preferentially drive the right hemisphere. The data support a model in which sounds are analyzed on two distinct timescales, 25-50 ms and 200-300 ms.


ALG2 : Science. 2005 Aug 5;309(5736):951-4.

Coupling between neuronal firing, field potentials, and FMRI in human auditory cortex.

Mukamel R, Gelbard H, Arieli A, Hasson U, Fried I, Malach R.

Functional magnetic resonance imaging (fMRI) is an important tool for investigating human brain function, but the relationship between the hemodynamically based fMRI signals in the human brain and the underlying neuronal activity is unclear. We recorded single unit activity and local field potentials in auditory cortex of two neurosurgical patients and compared them with the fMRI signals of 11 healthy subjects during presentation of an identical movie segment. The predicted fMRI signals derived from single units and the measured fMRI signals from auditory cortex showed a highly significant correlation (r = 0.75, P < 10(-47)). Thus, fMRI signals can provide a reliable measure of the firing rate of human cortical neurons.


ALG3 : Nat Neurosci. 2005 Sep;8(9):1241-7.

Structural and functional asymmetry of lateral Heschl's gyrus reflects pitch perception preference.

Schneider P, Sluming V, Roberts N, Scherg M, Goebel R, Specht HJ, Dosch HG, Bleeck S, Stippich C, Rupp A.

The relative pitch of harmonic complex sounds, such as instrumental sounds, may be perceived by decoding either the fundamental pitch (f0) or the spectral pitch (fSP) of the stimuli. We classified a large cohort of 420 subjects including symphony orchestra musicians to be either f0 or fSP listeners, depending on the dominant perceptual mode. In a subgroup of 87 subjects, MRI (magnetic resonance imaging) and magnetoencephalography studies demonstrated a strong neural basis for both types of pitch perception irrespective of musical aptitude. Compared with f0 listeners, fSP listeners possessed a pronounced rightward, rather than leftward, asymmetry of gray matter volume and P50m activity within the pitch-sensitive lateral Heschl's gyrus. Our data link relative hemispheric lateralization with perceptual stimulus properties, whereas the absolute size of the Heschl's gyrus depends on musical aptitude.


ALG4 : PLoS Biol. 2006 Jun;4(7):e215.

Functional imaging reveals numerous fields in the monkey auditory cortex.

Petkov CI, Kayser C, Augath M, Logothetis NK.

Anatomical studies propose that the primate auditory cortex contains more fields than have actually been functionally confirmed or described. Spatially resolved functional magnetic resonance imaging (fMRI) with carefully designed acoustical stimulation could be ideally suited to extend our understanding of the processing within these fields. However, after numerous experiments in humans, many auditory fields remain poorly characterized. Imaging the macaque monkey is of particular interest as these species have a richer set of anatomical and neurophysiological data to clarify the source of the imaged activity. We functionally mapped the auditory cortex of behaving and of anesthetized macaque monkeys with high resolution fMRI. By optimizing our imaging and stimulation procedures, we obtained robust activity throughout auditory cortex using tonal and band-passed noise sounds. Then, by varying the frequency content of the sounds, spatially specific activity patterns were observed over this region. As a result, the activity patterns could be assigned to many auditory cortical fields, including those whose functional properties were previously undescribed. The results provide an extensive functional tessellation of the macaque auditory cortex and suggest that 11 fields contain neurons tuned for the frequency of sounds. This study provides functional support for a model where three fields in primary auditory cortex are surrounded by eight neighboring 'belt’ fields in non-primary auditory cortex. The findings can now guide neurophysiological recordings in the monkey to expand our understanding of the processing within these fields. Additionally, this work will improve fMRI investigations of the human auditory cortex.


ALG6 : Proc Natl Acad Sci U S A. 2005 Jan 25;102(4):1181-6.

Visual speech speeds up the neural processing of auditory speech.

van Wassenhove V, Grant KW, Poeppel D.

Synchronous presentation of stimuli to the auditory and visual systems can modify the formation of a percept in either modality. For example, perception of auditory speech is improved when the speaker's facial articulatory movements are visible. Neural convergence onto multisensory sites exhibiting supra-additivity has been proposed as the principal mechanism for integration. Recent findings, however, have suggested that putative sensory-specific cortices are responsive to inputs presented through a different modality. Consequently, when and where audiovisual representations emerge remain unsettled. In combined psychophysical and electroencephalography experiments we show that visual speech speeds up the cortical processing of auditory signals early (within 100 ms of signal onset). The auditory-visual interaction is reflected as an articulator-specific temporal facilitation (as well as a nonspecific amplitude reduction). The latency facilitation systematically depends on the degree to which the visual signal predicts possible auditory targets. The observed auditory-visual data support the view that there exist abstract internal representations that constrain the analysis of subsequent speech inputs. This is evidence for the existence of an "analysis-by-synthesis" mechanism in auditory-visual speech perception.


AdC1 : Biology Letters. 2006 published online:1-4.

Correlated evolution between hearing sensitivity and social calls in bats

Bohn KM, Moss CF, Wilkinson GS

Echolocating bats are auditory specialists, with exquisite hearing that spans several octaves. In the ultrasonic range, bat audiograms typically show highest sensitivity in the spectral region of their species-specific echolocation calls. Welldeveloped hearing in the audible range has been commonly attributed to a need to detect sounds produced by prey. However, bat pups often emit isolation calls with low-frequency components that facilitate mother-young reunions. In this study, we examine whether low-frequency hearing in bats exhibits correlated evolution with (i) body size; (ii) high-frequency hearing sensitivity or (iii) pup isolation call frequency. Using published audiograms, we found that low-frequency hearing sensitivity is not dependent on body size but is related to high-frequency hearing. After controlling for high-frequency hearing, we found that low-frequency hearing exhibits correlated evolution with isolation call frequency. We infer that detection and discrimination of isolation calls have favoured enhanced lowfrequency hearing because accurate parental investment is critical: bats have low reproductive rates, non-volant altricial young and must often identify their pups within large crèches.


AdC2 : Nat Neurosci. 2006 Dec;9(12):1474-1476.

Cochlear efferent feedback balances interaural sensitivity.

Darrow KN, Maison SF, Liberman MC.

Neurons in the lateral superior olive (LSO) compute sound location based on differences in interaural intensity, coded in ascending signals from the two cochleas. Unilateral destruction of the neuronal feedback from the LSO to the cochlea, the lateral olivocochlear efferents, disrupted the normal interaural correlation in response amplitudes to sounds of equal intensity. Thus, lateral olivocochlear feedback maintains the binaural balance in neural excitability required for accurate localization of sounds in space.


AdC3 : Nat Neurosci. 2005 Dec;8(12):1684-9.

Neural population coding of sound level adapts to stimulus statistics.

Dean I, Harper NS, McAlpine D.

Mammals can hear sounds extending over a vast range of sound levels with remarkable accuracy. How auditory neurons code sound level over such a range is unclear; firing rates of individual neurons increase with sound level over only a very limited portion of the full range of hearing. We show that neurons in the auditory midbrain of the guinea pig adjust their responses to the mean, variance and more complex statistics of sound level distributions. We demonstrate that these adjustments improve the accuracy of the neural population code close to the region of most commonly occurring sound levels. This extends the range of sound levels that can be accurately encoded, fine-tuning hearing to the local acoustic environment.


AdC4 : Neuron. 2005 Nov 3;48(3):479-88.

Reliability and representational bandwidth in the auditory cortex.

DeWeese MR, Hromadka T, Zador AM.

It is unclear why there are so many more neurons in sensory cortex than in the sensory periphery. One possibility is that these "extra" neurons are used to overcome cortical noise and faithfully represent the acoustic stimulus. Another possibility is that even after overcoming cortical noise, there is "excess representational bandwidth" available and that this bandwidth is used to represent conjunctions of auditory and nonauditory information for computation. Here, we discuss recent data about neuronal reliability in auditory cortex showing that cortical noise may not be as high as was previously believed. Although at present, the data suggest that auditory cortex neurons can be more reliable than those in the visual cortex, we speculate that the principles governing cortical computation are universal and that visual and other cortical areas can also exploit strategies based on similarly high-fidelity activity.


AdC5 : J Acoust Soc Am Dec 2006 120:3907-15

Individual differences in the sensitivity to pitch direction.

Semal C, Demany L.

It is commonly assumed that one can always assign a direction -upward or downward- to a percept of pitch change. The present study shows that this is true for some, but not all, listeners. Frequency difference limens (FDLs, in cents) for pure tones roved in frequency were measured in two conditions. In one condition, the task was to detect frequency changes; in the other condition, the task was to identify the direction of frequency changes. For three listeners, the identification FDL was about 1.5 times smaller than the detection FDL, as predicted (counterintuitively) by signal detection theory under the assumption that performance in the two conditions was limited by one and the same internal noise. For three other listeners, however, the identification FDL was much larger than the detection FDL. The latter listeners had relatively high detection FDLs. They had no difficulty in identifying the direction of just-detectable changes in intensity, or in the frequency of amplitude modulation. Their difficulty in perceiving the direction of small frequency/pitch changes showed up not only when the task required absolute judgments of direction, but also when the directions of two successive frequency changes had to be judged as identical or different


BT1 : Proc Natl Acad Sci U S A. 2005 Aug 30;102(35):12639-43.

Tuning in to musical rhythms: infants learn more readily than adults.

Hannon EE, Trehub SE.

Domain-general tuning processes may guide the acquisition of perceptual knowledge in infancy. Here, we demonstrate that 12-month-old infants show an adult-like, culture-specific pattern of responding to musical rhythms, in contrast to the culture-general responding that is evident at 6 months of age. Nevertheless, brief exposure to foreign music enables 12-month-olds, but not adults, to perceive rhythmic distinctions in foreign musical contexts. These findings may indicate a sensitive period early in life for acquiring rhythm in particular or socially and biologically important structures more generally.


BT2 : J Cogn Neurosci. 2005 Oct;17(10):1565-77.

Interaction between syntax processing in language and in music: an ERP Study.

Koelsch S, Gunter TC, Wittfoth M, Sammler D.

The present study investigated simultaneous processing of language and music using visually presented sentences and auditorily presented chord sequences. Music-syntactically regular and irregular chord functions were presented synchronously with syntactically correct or incorrect words, or with words that had either a high or a low semantic cloze probability. Music-syntactically irregular chords elicited an early right anterior negativity (ERAN). Syntactically incorrect words elicited a left anterior negativity (LAN). The LAN was clearly reduced when words were presented simultaneously with music-syntactically irregular chord functions. Processing of high and low cloze-probability words as indexed by the N400 was not affected by the presentation of irregular chord functions. In a control experiment, the LAN was not affected by physically deviant tones that elicited a mismatch negativity (MMN). Results demonstrate that processing of musical syntax (as reflected in the ERAN) interacts with the processing of linguistic syntax (as reflected in the LAN), and that this interaction is not due to a general effect of deviance-related negativities that precede an LAN. Findings thus indicate a strong overlap of neural resources involved in the processing of syntax in language and music.


BT3 : Cognition. 2000 Jul 14;76(1):13-58.

Cross-cultural music cognition: cognitive methodology applied to North Sami yoiks.

Krumhansl CL, Toivanen P, Eerola T, Toiviainen P, Jarvinen T, Louhivuori J.

This article is a study of melodic expectancy in North Sami yoiks, a style of music quite distinct from Western tonal music. Three different approaches were taken. The first approach was a statistical style analysis of tones in a representative corpus of 18 yoiks. The analysis determined the relative frequencies of tone onsets and two- and three-tone transitions. It also identified style characteristics, such as pentatonic orientation, the presence of two reference pitches, the frequency of large consonant intervals, and a relatively large set of possible melodic continuations. The second approach was a behavioral experiment in which listeners made judgments about melodic continuations. Three groups of listeners participated. One group was from the Sami culture, the second group consisted of Finnish music students who had learned some yoiks, and the third group consisted of Western musicians unfamiliar with yoiks. Expertise was associated with stronger veridical expectations (for the correct next tone) than schematic expectations (based on general style characteristics). Familiarity with the particular yoiks was found to compensate for lack of experience with the musical culture. The third approach simulated melodic expectancy with neural network models of the self-organizing map (SOM) type (Kohonen, T. (1997). Self-organizing maps (2nd ed.). Berlin: Springer). One model was trained on the excerpts of yoiks used in the behavioral experiment including the correct continuation tone, while another was trained with a set of Finnish folk songs and Lutheran hymns. The convergence of the three approaches showed that both listeners and the SOM model are influenced by the statistical distributions of tones and tone sequences. The listeners and SOM models also provided evidence supporting a core set of psychological principles underlying melody formation whose relative weights appear to differ across musical styles.


BT4 : Psychophysiology. 2004 May;41(3):341-9.

The music of speech: music training facilitates pitch processing in both music and language.

Schon D, Magne C, Besson M.

The main aim of the present experiment was to determine whether extensive musical training facilitates pitch contour processing not only in music but also in language. We used a parametric manipulation of final notes' or words' fundamental frequency (F0), and we recorded behavioral and electrophysiological data to examine the precise time course of pitch processing. We compared professional musicians and nonmusicians. Results revealed that within both domains, musicians detected weak F0 manipulations better than nonmusicians. Moreover, F0 manipulations within both music and language elicited similar variations in brain electrical potentials, with overall shorter onset latency for musicians than for nonmusicians. Finally, the scalp distribution of an early negativity in the linguistic task varied with musical expertise, being largest over temporal sites bilaterally for musicians and largest centrally and over left temporal sites for nonmusicians. These results are taken as evidence that extensive musical training influences the perception of pitch contour in spoken language.


CL1 : Eur J Neurosci. 2006 Oct;24(7):2003-10.

The effects of intense sound exposure on phase locking in the chick (Gallus domesticus) cochlear nerve.

Furman AC, Avissar M, Saunders JC.

Little is known about changes that occur to phase locking in the auditory nerve following exposure to intense and damaging levels of sound. The present study evaluated synchronization in the discharge patterns of cochlear nerve units collected from two groups of young chicks (Gallus domesticus), one shortly after removal from an exposure to a 120-dB, 900-Hz pure tone for 48 h and the other from a group of non-exposed control animals. Spontaneous activity, the characteristic frequency (CF), CF threshold and a phaselocked peri-stimulus time histogram were obtained for every unit in each group. Vector strength and temporal dispersion were calculated from these peri-stimulus time histograms, and plotted against the unit’s CF. All parameters of unit responses were then compared between control and exposed units. The results in exposed units revealed that CF thresholds were elevated by 30-35 dB whereas spontaneous activity declined by 24%. In both control and exposed units a high degree of synchronization was observed in the low frequencies. The level of synchronization above approximately 0.5 kHz then systematically declined. The vector strengths in units recorded shortly after removal from the exposure were identical to those seen in control chicks. The deterioration in discharge activity of exposed units, seen in CF threshold and spontaneous activity, contrasted with the total absence of any overstimulation effect on synchronization. This suggested that synchronization arises from mechanisms unscathed by the acoustic trauma induced by the exposure.


CL2 : J Acoust Soc Am. 2003 Feb;113(2):961-8.

Understanding speech in modulated interference: cochlear implant users and normal-hearing listeners.

Nelson PB, Jin SH, Carney AE, Nelson DA.

Many competing noises in real environments are modulated or fluctuating in level. Listeners with normal hearing are able to take advantage of temporal gaps in fluctuating maskers. Listeners with sensorineural hearing loss show less benefit from modulated maskers. Cochlear implant users may be more adversely affected by modulated maskers because of their limited spectral resolution and by their reliance on envelope-based signal-processing strategies of implant processors. The current study evaluated cochlear implant users' ability to understand sentences in the presence of modulated speech-shaped noise. Normal-hearing listeners served as a comparison group. Listeners repeated IEEE sentences in quiet, steady noise, and modulated noise maskers. Maskers were presented at varying signal-to-noise ratios (SNRs) at six modulation rates varying from 1 to 32 Hz. Results suggested that normal-hearing listeners obtain significant release from masking from modulated maskers, especially at 8-Hz masker modulation frequency. In contrast, cochlear implant users experience very little release from masking from modulated maskers. The data suggest, in fact, that they may show negative effects of modulated maskers at syllabic modulation rates (2-4 Hz). Similar patterns of results were obtained from implant listeners using three different devices with different speech-processor strategies. The lack of release from masking occurs in implant listeners independent of their device characteristics, and may be attributable to the nature of implant processing strategies and/or the lack of spectral detail in processed stimuli.


CL3 : J Acoust Soc Am. 2005 Oct;118(4):2519-26.

Consequences of cochlear damage for the detection of interaural phase differences.

Lacher-Fougere S, Demany L.

Thresholds for detecting interaural phase differences (IPDs) in sinusoidally amplitude-modulated pure tones were measured in seven normal-hearing listeners and nine listeners with bilaterally symmetric hearing losses of cochlear origin. The IPDs were imposed either on the carrier signal alone-not the amplitude modulation-or vice versa. The carrier frequency was 250, 500, or 1000 Hz, the modulation frequency 20 or 50 Hz, and the sound pressure level was fixed at 75 dB. A three-interval two-alternative forced choice paradigm was used. For each type of IPD (carrier or modulation), thresholds were on average higher for the hearing-impaired than for the normal listeners. However, the impaired listeners' detection deficit was markedly larger for carrier IPDs than for modulation IPDs. This was not predictable from the effect of hearing loss on the sensation level of the stimuli since, for normal listeners, large reductions of sensation level appeared to be more deleterious to the detection of modulation IPDs than to the detection of carrier IPDs. The results support the idea that one consequence of cochlear damage is a deterioration in the perceptual sensitivity to the temporal fine structure of sounds.


CL4 : J Acoust Soc Am. 1998 Jan;103(1):577-87.

Speech reception thresholds in noise with and without spectral and temporal dips for hearing-impaired and normally hearing people.

Peters RW, Moore BC, Baer T.

People with cochlear hearing loss often have considerable difficulty in understanding speech in the presence of background sounds. In this paper the relative importance of spectral and temporal dips in the background sounds is quantified by varying the degree to which they contain such dips. Speech reception thresholds in a 65-dB SPL noise were measured for four groups of subjects: (a) young with normal hearing; (b) elderly with near-normal hearing; (c) young with moderate to severe cochlear hearing loss; and (d) elderly with moderate to severe cochlear hearing loss. The results indicate that both spectral and temporal dips are important. In a background that contained both spectral and temporal dips, groups (c) and (d) performed much more poorly than group (a). The signal-to-background ratio required for 50% intelligibility was about 19 dB higher for group (d) than for group (a). Young hearing-impaired subjects showed a slightly smaller deficit, but still a substantial one. Linear amplification combined with appropriate frequency-response shaping (NAL amplification), as would be provided by a well-fitted "conventional" hearing aid, only partially compensated for these deficits. For example, group (d) still required a speech-to-background ratio that was 15 dB higher than for group (a). Calculations of the articulation index indicated that NAL amplification did not restore audibility of the whole of the speech spectrum when the speech-to-background ratio was low. For unamplified stimuli, the SRTs in background sounds were highly correlated with absolute thresholds, but not with age. For stimuli with NAL amplification, the correlations of SRTs with absolute thresholds were lower, but SRTs in backgrounds with spectral and/or temporal dips were significantly correlated with age. It is proposed that noise with spectral and temporal dips may be especially useful in evaluating possible benefits of multi-channel compression.


CL5 : Proc Natl Acad Sci U S A. 2005 Feb 15;102(7):2293-8.

Speech recognition with amplitude and frequency modulations.

Zeng FG, Nie K, Stickney GS, Kong YY, Vongphoe M, Bhargave A, Wei C, Cao K.

Amplitude modulation (AM) and frequency modulation (FM) are commonly used in communication, but their relative contributions to speech recognition have not been fully explored. To bridge this gap, we derived slowly varying AM and FM from speech sounds and conducted listening tests using stimuli with different modulations in normal-hearing and cochlear-implant subjects. We found that although AM from a limited number of spectral bands may be sufficient for speech recognition in quiet, FM significantly enhances speech recognition in noise, as well as speaker and tone recognition. Additional speech reception threshold measures revealed that FM is particularly critical for speech recognition with a competing voice and is independent of spectral resolution and similarity. These results suggest that AM and FM provide independent yet complementary contributions to support robust speech recognition under realistic listening situations. Encoding FM may improve auditory scene analysis, cochlear-implant, and audiocoding performance.


DP1 : J Acoust Soc Am. 2005 Feb;117(2):833-41.

On the binding of successive sounds: perceiving shifts in nonperceived pitches.

Demany L, Ramos C.

It is difficult to hear out individually the components of a "chord" of equal-amplitude pure tones with synchronous onsets and offsets. In the present study, this was confirmed using 300-ms random (inharmonic) chords with components at least 1/2 octave apart. Following each chord, after a variable silent delay, listeners were presented with a single pure tone which was either identical to one component of the chord or halfway in frequency between two components. These two types of sequence could not be reliably discriminated from each other. However, it was also found that if the single tone following the chord was instead slightly (e.g., 1/12 octave) lower or higher in frequency than one of its components, the same listeners were sensitive to this relation. They could perceive a pitch shift in the corresponding direction. Thus, it is possible to perceive a shift in a nonperceived frequency/pitch. This paradoxical phenomenon provides psychophysical evidence for the existence of automatic "frequency-shift detectors" in the human auditory system. The data reported here suggest that such detectors operate at an early stage of auditory scene analysis but can be activated by a pair of sounds separated by a few seconds.


DP2 : Curr Biol. 2005 Jun 21;15(12):1108-13.

Directed attention eliminates 'change deafness' in complex auditory scenes.

Eramudugolla R, Irvine DR, McAnally KI, Martin RL, Mattingley JB.

In natural environments that contain multiple sound sources, acoustic energy arising from the different sources sums to produce a single complex waveform at each of the listener's ears. The auditory system must segregate this waveform into distinct streams to permit identification of the objects from which the signals emanate [1]. Although the processes involved in stream segregation are now reasonably well understood [1, 2 and 3], little is known about the nature of our perception of complex auditory scenes. Here, we examined complex scene perception by having listeners detect a discrete change to an auditory scene comprising multiple concurrent naturalistic sounds. We found that listeners were remarkably poor at detecting the disappearance of an individual auditory object when listening to scenes containing more than four objects, but they performed near perfectly when their attention was directed to the identity of a potential change. In the absence of directed attention, this "change deafness" [4] was greater for objects arising from a common location in space than for objects separated in azimuth. Change deafness was also observed for changes in object location, suggesting that it may reflect a general effect of the dependence of human auditory perception on attention.


DP3 : Neuron. 2005 Oct 6;48(1):139-48.

Perceptual organization of tone sequences in the auditory cortex of awake macaques.

Micheyl C, Tian B, Carlyon RP, Rauschecker JP.

Acoustic sequences such as speech and music are generally perceived as coherent auditory "streams," which can be individually attended to and followed over time. Although the psychophysical stimulus parameters governing this "auditory streaming" are well established, the brain mechanisms underlying the formation of auditory streams remain largely unknown. In particular, an essential feature of the phenomenon, which corresponds to the fact that the segregation of sounds into streams typically takes several seconds to build up, remains unexplained. Here, we show that this and other major features of auditory-stream formation measured in humans using alternating-tone sequences can be quantitatively accounted for based on single-unit responses recorded in the primary auditory cortex (A1) of awake rhesus monkeys listening to the same sound sequences.


DP4 : Proc Natl Acad Sci U S A. 2004 Feb 3;101(5):1421-5.

Correct tonotopic representation is necessary for complex pitch perception.

Oxenham AJ, Bernstein JG, Penagos H.

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.


DP5 : Hear Res. 2006 Sep;219(1-2):36-47.

Influence of musical and psychoacoustical training on pitch discrimination.

Micheyl C, Delhommeau K, Perrot X, Oxenham AJ.

This study compared the influence of musical and psychoacoustical training on auditory pitch discrimination abilities. In a first experiment, pitch discrimination thresholds for pure and complex tones were measured in 30 classical musicians and 30 non-musicians, none of whom had prior psychoacoustical training. The non-musicians’ mean thresholds were more than six times larger than those of the classical musicians initially, and still about four times larger after 2 h of training using an adaptive two-interval forced-choice procedure; this difference is two to three times larger than suggested by previous studies. The musicians’ thresholds were close to those measured in earlier psychoacoustical studies using highly trained listeners, and showed little improvement with training; this suggests that classical musical training can lead to optimal or nearly optimal pitch discrimination performance. A second experiment was performed to determine how much additional training was required for the non-musicians to obtain thresholds as low as those of the classical musicians from experiment 1. Eight new non-musicians with no prior training practiced the frequency discrimination task for a total of 14 h. It took between 4 and 8 h of training for their thresholds to become as small as those measured in the classical musicians from experiment 1. These findings supplement and qualify earlier data in the literature regarding the respective influence of musical and psychoacoustical training on pitch discrimination performance.


DP6 : Nat Neurosci. 2003 Apr;6(4):391-8.

Processing of low-probability sounds by cortical neurons.

Ulanovsky N, Las L, Nelken I.

The ability to detect rare auditory events can be critical for survival. We report here that neurons in cat primary auditory cortex (A1) responded more strongly to a rarely presented sound than to the same sound when it was common. For the rare stimuli, we used both frequency and amplitude deviants. Moreover, some A1 neurons showed hyperacuity for frequency deviants--a frequency resolution one order of magnitude better than receptive field widths in A1. In contrast, auditory thalamic neurons were insensitive to the probability of frequency deviants. These phenomena resulted from stimulus-specific adaptation in A1, which may be a single-neuron correlate of an extensively studied cortical potential--mismatch negativity--that is evoked by rare sounds. Our results thus indicate that A1 neurons, in addition to processing the acoustic features of sounds, may also be involved in sensory memory and novelty detection.


DP7 : .Ear Hear. 2004 Apr;25(2):173-85.

Music perception with temporal cues in acoustic and electric hearing.

Kong YY, Cruz R, Jones JA, Zeng FG.

OBJECTIVE: The first specific aim of the present study is to compare the ability of normal-hearing and cochlear implant listeners to use temporal cues in three music perception tasks: tempo discrimination, rhythmic pattern identification, and melody identification. The second aim is to identify the relative contribution of temporal and spectral cues to melody recognition in acoustic and electric hearing. DESIGN: Both normal-hearing and cochlear implant listeners participated in the experiments. Tempo discrimination was measured in a two-interval forced-choice procedure in which subjects were asked to choose the faster tempo at four standard tempo conditions (60, 80, 100, and 120 beats per minute). For rhythmic pattern identification, seven different rhythmic patterns were created and subjects were asked to read and choose the musical notation displayed on the screen that corresponded to the rhythmic pattern presented. Melody identification was evaluated with two sets of 12 familiar melodies. One set contained both rhythm and melody information (rhythm condition), whereas the other set contained only melody information (no-rhythm condition). Melody stimuli were also processed to extract the slowly varying temporal envelope from 1, 2, 4, 8, 16, 32, and 64 frequency bands, to create cochlear implant simulations. Subjects listened to a melody and had to respond by choosing one of the 12 names corresponding to the melodies displayed on a computer screen. RESULTS: In tempo discrimination, the cochlear implant listeners performed similarly to the normal-hearing listeners with rate discrimination difference limens obtained at 4-6 beats per minute. In rhythmic pattern identification, the cochlear implant listeners performed 5-25 percentage points poorer than the normal-hearing listeners. The normal-hearing listeners achieved perfect scores in melody identification with and without the rhythmic cues. However, the cochlear implant listeners performed significantly poorer than the normal-hearing listeners in both rhythm and no-rhythm conditions. The simulation results from normal-hearing listeners showed a relatively high level of performance for all numbers of frequency bands in the rhythm condition but required as many as 32 bands in the no-rhythm condition. CONCLUSIONS: Cochlear-implant listeners performed normally in tempo discrimination, but significantly poorer than normal-hearing listeners in rhythmic pattern identification and melody recognition. While both temporal (rhythmic) and spectral (pitch) cues contribute to melody recognition, cochlear-implant listeners mostly relied on the rhythmic cues for melody recognition. Without the rhythmic cues, high spectral resolution with as many as 32 bands was needed for melody recognition for normal-hearing listeners. This result indicates that the present cochlear implants provide sufficient spectral cues to support speech recognition in quiet, but they are not adequate to support music perception. Increasing the number of functional channels and improved encoding of the fine structure information are necessary to improve music perception for cochlear implant listeners.


JME1 : Proc Natl Acad Sci U S A. 2003 Feb 4;100(3):1405-8.

Suppression of cortical representation through backward conditioning.

Bao S, Chan VT, Zhang LI, Merzenich MM.

Temporal stimulus reinforcement sequences have been shown to determine the directions of synaptic plasticity and behavioral learning. Here, we examined whether they also control the direction of cortical reorganization. Pairing ventral tegmental area stimulation with a sound in a backward conditioning paradigm specifically reduced representations of the paired sound in the primary auditory cortex (AI). This temporal sequence-dependent bidirectional cortical plasticity modulated by dopamine release hypothetically serves to prevent the over-representation of frequently occurring stimuli resulting from their random pairing with unrelated rewards.


JME2 : J Neurosci. 2005 Mar 9;25(10):2490-503.

Plasticity in primary auditory cortex of monkeys with altered vocal production.

Cheung SW, Nagarajan SS, Schreiner CE, Bedenbaugh PH, Wong A.

Response properties of primary auditory cortical neurons in the adult common marmoset monkey (Callithrix jacchus) were modified by extensive exposure to altered vocalizations that were self-generated and rehearsed frequently. A laryngeal apparatus modification procedure permanently lowered the frequency content of the native twitter call, a complex communication vocalization consisting of a series of frequency modulation (FM) sweeps. Monkeys vocalized shortly after this procedure and maintained voicing efforts until physiological evaluation 5-15 months later. The altered twitter calls improved over time, with FM sweeps approaching but never reaching the normal spectral range. Neurons with characteristic frequencies <4.3 kHz that had been weakly activated by native twitter calls were recruited to encode self-uttered altered twitter vocalizations. These neurons showed a decrease in response magnitude and an increase in temporal dispersion of response timing to twitter call and parametric FM stimuli but a normal response profile to pure tone stimuli. Tonotopic maps in voice-modified monkeys were not distorted. These findings suggest a previously unrecognized form of cortical plasticity that is specific to higher-order processes involved in the discrimination of more complex sounds, such as species-specific vocalizations.


JME3 : Eur J Neurosci. 2006 Aug;24(3):857-66.

Neonatal nicotine exposure impairs nicotinic enhancement of central auditory processing and auditory learning in adult rats.

Liang K, Poytress BS, Chen Y, Leslie FM, Weinberger NM, Metherate R.

Children of women who smoke cigarettes during pregnancy display cognitive deficits in the auditory-verbal domain. Clinical studies have implicated developmental exposure to nicotine, the main psychoactive ingredient of tobacco, as a probable cause of subsequent auditory deficits. To test for a causal link, we have developed an animal model to determine how neonatal nicotine exposure affects adult auditory function. In adult control rats, nicotine administered systemically (0.7 mg/kg, s.c.) enhanced the sensitivity to sound of neural responses recorded in primary auditory cortex. The effect was strongest in cortical layers 3 and 4, where there is a dense concentration of nicotinic acetylcholine receptors (nAChRs) that has been hypothesized to regulate thalamocortical inputs. In support of the hypothesis, microinjection into layer 4 of the nonspecific nAChR antagonist mecamylamine (10 microM) strongly reduced sound-evoked responses. In contrast to the effects of acute nicotine and mecamylamine in adult control animals, neither drug was as effective in adult animals that had been treated with 5 days of chronic nicotine exposure (CNE) shortly after birth. Neonatal CNE also impaired performance on an auditory-cued active avoidance task, while having little effect on basic auditory or motor functions. Thus, neonatal CNE impairs nicotinic regulation of cortical function, and auditory learning, in the adult. Our results provide evidence that developmental nicotine exposure is responsible for auditory-cognitive deficits in the offspring of women who smoke during pregnancy, and suggest a potential underlying mechanism, namely diminished function of cortical nAChRs.


JME4 : Eur J Neurosci. 2006 Jun;23(11):3087-97.

Improved cortical entrainment to infant communication calls in mothers compared with virgin mice.

Liu RC, Linden JF, Schreiner CE.

There is a growing interest in the use of mice as a model system for species-specific communication. In particular, ultrasonic calls emitted by mouse pups communicate distress, and elicit a search and retrieval response from mothers. Behaviorally, mothers prefer and recognize these calls in two-alternative choice tests, in contrast to pup-naive females that do not have experience with pups. Here, we explored whether one particular acoustic feature that defines these calls-- the repetition rate of calls within a bout-- is represented differently in the auditory cortex of these two animal groups. Multiunit recordings in anesthetized CBA/CaJ mice revealed that: (i) neural entrainment to repeated stimuli extended up to the natural pup call repetition rate (5 Hz) in mothers; but (ii) neurons in naive females followed repeated stimuli well only at slower repetition rates; and (iii) entrained responses to repeated pup calls were less sensitive to natural pup call variability in mothers than in pup-naive females. In the broader context, our data suggest that auditory cortical responses to communication sounds are plastic, and that communicative significance is correlated with an improved cortical representation.


JME5 : Proc Natl Acad Sci U S A. 2004 Nov 16;101(46):16351-6.

Associative learning shapes the neural code for stimulus magnitude in primary auditory cortex.

Polley DB, Heiser MA, Blake DT, Schreiner CE, Merzenich MM.

Since the dawn of experimental psychology, researchers have sought an understanding of the fundamental relationship between the amplitude of sensory stimuli and the magnitudes of their perceptual representations. Contemporary theories support the view that magnitude is encoded by a linear increase in firing rate established in the primary afferent pathways. In the present study, we have investigated sound intensity coding in the rat primary auditory cortex (AI) and describe its plasticity by following paired stimulus reinforcement and instrumental conditioning paradigms. In trained animals, population-response strengths in AI became more strongly nonlinear with increasing stimulus intensity. Individual AI responses became selective to more restricted ranges of sound intensities and, as a population, represented a broader range of preferred sound levels. These experiments demonstrate that the representation of stimulus magnitude can be powerfully reshaped by associative learning processes and suggest that the code for sound intensity within AI can be derived from intensity-tuned neurons that change, rather than simply increase, their firing rates in proportion to increases in sound intensity.


JME6 : J Neurosci. 2006 May 3;26(18):4785-95.

Plasticity of temporal pattern codes for vocalization stimuli in primary auditory cortex.

Schnupp JW, Hall TM, Kokelaar RF, Ahmed B.

It has been suggested that "call-selective" neurons may play an important role in the encoding of vocalizations in primary auditory cortex (A1). For example, marmoset A1 neurons often respond more vigorously to natural than to time-reversed twitter calls, although the spectral energy distribution in the natural and time-reversed signals is the same. Neurons recorded in cat A1, in contrast, showed no such selectivity for natural marmoset calls. To investigate whether call selectivity in A1 can arise purely as a result of auditory experience, we recorded responses to marmoset calls in A1 of naive ferrets, as well as in ferrets that had been trained to recognize these natural marmoset calls. We found that training did not induce call selectivity for the trained vocalizations in A1. However, although ferret A1 neurons were not call selective, they efficiently represented the vocalizations through temporal pattern codes, and trained animals recognized marmoset twitters with a high degree of accuracy. These temporal patterns needed to be analyzed at timescales of 10-50 ms to ensure efficient decoding. Training led to a substantial increase in the amount of information transmitted by these temporal discharge patterns, but the fundamental nature of the temporal pattern code remained unaltered. These results emphasize the importance of temporal discharge patterns and cast doubt on the functional significance of call-selective neurons in the processing of animal communication sounds at the level of A1.