Pitch Perception

Music and speech are very cognitively demanding auditory phenomena generally attributed to cortical rather than subcortical circuitry. We examined brainstem encoding of linguistic pitch and found that musicians show more robust and faithful encoding compared with nonmusicians. These results not only implicate a common subcortical manifestation for two presumed cortical functions, but also a possible reciprocity of corticofugal speech and music tuning, providing neurophysiological explanations for musicians' higher language-learning ability.

Neuronally plausible, generative or forward models are essential for understanding how event-related fields (ERFs) and potentials (ERPs) are generated. In this paper we present a new approach to modeling eventrelated responses measured with EEG or MEG. This approach uses a biologically informed model to make inferences about the underlying neuronal networks generating responses. The approach can be regarded as a neurobiologically constrained source reconstruction scheme, in which the parameters of the reconstruction have an explicit neuronal interpretation. Specifically, these parameters encode, among other things, the coupling among sources and how that coupling depends upon stimulus attributes or experimental context. The basic idea is to supplement conventional electromagnetic forward models, of how sources are expressed in measurement space, with a model of how source activity is generated by neuronal dynamics.

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 ...

Song imitation in birds provides good material for studying the basic biology of vocal learning. Techniques were developed for inducing the rapid onset of song imitation in young zebra finches and for tracking trajectories of vocal change over a 7-week period until a match to a model song was achieved. Exposure to a model song induced the prompt generation of repeated structured sounds ( prototypes) followed by a slow transition from repetitive to serial delivery of syllables. Tracking this transition revealed two phenomena: (i) Imitations of dissimilar sounds can emerge from successive renditions of the same prototype, and (ii) developmental trajectories for some sounds followed paths of increasing acoustic mismatch until an abrupt correction occurred by period doubling. These dynamics are likely to reflect underlying neural and articulatory constraints on the production and imitation of sounds.

Auditory evoked magnetic fields of the human brain were recorded with a four-channel 1st order gradiometer. Pitch deviance in a sequence of repetitive tone pips elicited magnetic evoked-response changes with a topography suggesting that a neuronal mismatch process to the deviant tones activates the primary auditory cortex. 0304-3940/84/$ 03.00

Objectives/Hypothesis: This study documents the importance of preserving residual low-frequency acoustic hearing as those with more residual hearing are selected for cochlear implantation. Surgical strategies used for hearing preservation with a short hybrid cochlear implant are outlined. The benefits of preserved residual low-frequency hearing, improved word understanding in noise, and music appreciation are described. Study Design: Multicenter, prospective, single-subject design. Methods: Records were reviewed of 21 individuals participating in an Food and Drug Administration (FDA) feasibility clinical trial who have received an Iowa/Nucleus 10 mm electrode. A second group of subjects receiving implants at the University of Iowa that have used the 10 mm device between 2 years and 6 months were also reviewed. Outcome measures included standardized tests of monosyllabic word understanding, spondees in noise, and common melody recognition. Results: Lowfrequency hearing was maintained in all individuals immediately postoperative. One subject lost hearing at 2.5 months postoperative after a viral infection. The group has averaged a loss of ؊9 dB low-frequency acoustic hearing between 125 and 1,000 Hz. Monosyllabic word understanding scores at 6 months for a group being followed for an FDA clinical trial using the implant plus hearing aids was 69% correct. For the long-term group receiving implants at Iowa, monosyllabic word understanding in those who have used the device between 6 months and 2 years is 79%. Other important findings include improved recognition of speech in noise (9 dB improvement) as compared with standard cochlear implant recipients who were matched for speech recognition in quiet and near normal recognition of common melodies. Conclusion: The surgical strategies outlined have been successful in preservation of low-frequency hearing in 96% of individuals. Combined electrical and acoustical speech processing has enabled this group of volunteers to gain improved word understanding as compared with their preoperative hearing with bilateral hearing aids and a group of individuals receiving a standard cochlear implant with similar experience with their device. The improvement of speech in noise and melody recognition is attributed to the ability to distinguish fine pitch differences as the result of preserved residual low-frequency acoustic hearing. Preservation of low-frequency acoustic hearing is important for improving speech in noise and music appreciation for the hearing impaired, both of which are important in real-life situations.

Spectro-temporal response field characterization with dynamic ripples in ferret primary auditory cortex. J Neurophysiol 85: 1220 -1234, 2001. To understand the neural representation of broadband, dynamic sounds in primary auditory cortex (AI), we characterize responses using the spectro-temporal response field (STRF). The STRF describes, predicts, and fully characterizes the linear dynamics of neurons in response to sounds with rich spectrotemporal envelopes. It is computed from the responses to elementary "ripples," a family of sounds with drifting sinusoidal spectral envelopes. The collection of responses to all elementary ripples is the spectro-temporal transfer function. The complex spectro-temporal envelope of any broadband, dynamic sound can expressed as the linear sum of individual ripples. Previous experiments using ripples with downward drifting spectra suggested that the transfer function is separable, i.e., it is reducible into a product of purely temporal and purely spectral functions. Here we measure the responses to upward and downward drifting ripples, assuming reparability within each direction, to determine if the total bidirectional transfer function is fully separable. In general, the combined transfer function for two directions is not symmetric, and hence units in AI are not, in general, fully separable. Consequently, many AI units have complex response properties such as sensitivity to direction of motion, though most inseparable units are not strongly directionally selective. We show that for most neurons, the lack of full separability stems from differences between the upward and downward spectral cross-sections but not from the temporal cross-sections; this places strong constraints on the neural inputs of these AI units.

Visual and auditory motion information can be used together to provide complementary information about the movement of objects. To investigate the neural substrates of such cross-modal integration, functional magnetic resonance imaging was used to assess brain activation while subjects performed separate visual and auditory motion discrimination tasks. Areas of unimodal activation included the primary and/or early sensory cortex for each modality plus additional sites extending toward parietal cortex. Areas conjointly activated by both tasks included lateral parietal cortex, lateral frontal cortex, anterior midline and anterior insular cortex. The parietal site encompassed distinct, but partially overlapping, zones of activation in or near the intraparietal sulcus (IPS). A subsequent task requiring an explicit cross-modal speed comparison revealed several foci of enhanced activity relative to the unimodal tasks. These included the IPS, anterior midline, and anterior insula but not frontal cortex. During the unimodal auditory motion task, portions of the dorsal visual motion system showed signals depressed below resting baseline. Thus, interactions between the two systems involved either enhancement or suppression depending on the stimuli present and the nature of the perceptual task. Together, these results identify human cortical regions involved in polysensory integration and the attentional selection of cross-modal motion information.

The responses of neuronal clusters to amplitude-modulated tones were studied in five auditory cortical fields of the ancsthetiLed cat: the primary auditory field (Al). second auditory field (AII), anterior auditory field (AAF). posterior auditory field (PAF) and the ventro-posterior auditory field (VPAF). Modulation transfer functions (MTFs) for amplitude-modulated tones were obtained at 172 cortical locations. MTFs were constructed by measuring firing rate (rate-MTFs) and response s~nchr~nizatioll (synchr~~nizati(~n-MTFs) to sinusoidal and rectangular waveform modulation of CF-tones, The MTFs were characterized by their .best-rn[~duiati(~n frequency' (BMF) and a measure of their quality or 'sharpness' (Q z,ie). These characteristics were compared for the five fields. Rate and synchronization MTFs for sinusoidal and rectangular modulation produced similar estimates of BMF and Qz,,t~, Comparison of averaged BMFs between the cortical fields revealed relatively high BMFs in AAF (mean: 31.1 Hz for synchronization to sinusoidal AM) and moderately high BMFs in AI (14.2 Hz) whereas BMFs encountered in All. VPAF and PAF were generally low (7.0, 5.2. and 6.8 Hz). The MTFs were relatively broadly tuned (low Qzda) in AAF and sharper in a low modulation group containing AII. PAF and VPAF. The ventro-posterior field was the most sensitive to changes m the modujati(~n waveform. We conclude that there are significant differences between auditory cortical fields with respect to their temporal response characteristics and that the assessment of these response characteristics reveals important aspects of the functional significance of auditory cortical fields for the coding and representation of complex sounds Auditory cortex: Amplitude modulation: Cat

The cerebral representation of the temporal envelope of sounds was studied in five normal-hearing subjects using functional magnetic resonance imaging. The stimuli were white noise, sinusoidally amplitude-modulated at frequencies ranging from 4 to 256 Hz. This range includes low AM frequencies (up to 32 Hz) essential for the perception of the manner of articulation and syllabic rate, and high AM frequencies (above 64 Hz) essential for the perception of voicing and prosody. The right lower brainstem (superior olivary complex), the right inferior colliculus, the left medial geniculate body, Heschl's gyrus, the superior temporal gyrus, the superior temporal sulcus, and the inferior parietal lobule were specifically responsive to AM. Global tuning curves in these regions suggest that the human auditory system is organized as a hierarchical filter bank, each processing level responding preferentially to a given AM frequency, 256 Hz for the lower brainstem, 32-256 Hz for the inferior ...

Experience-dependent plasticity during development results in the emergence of highly adapted representations of the external world in the adult brain. Previous studies have convincingly shown that the primary auditory cortex (A1) of the rat possesses a postnatal period of sensory input-driven plasticity but its precise timing (onset, duration, end) has not been defined. In the present study, we examined the effects of pure-tone exposure on the auditory cortex of developing rat pups at different postnatal ages with a high temporal resolution. We found that pure-tone exposure resulted in profound, persistent alterations in sound representations in A1 only if the exposure occurred during a brief period extending from postnatal day 11 (P11) to P13. We also found that postnatal sound exposure in this epoch led to striking alterations in the cortical representation of sound intensity.

Reading acquisition involves two fundamental processes: learning to decode the print (i.e., visually recognize the words), and comprehending the meanings of the print (Gough & Tunmer, 1986; Hoover & Gough, 1990). Comprehension skill is acquired in the course of learning to speak, and reading comprehension processes are very similar to listening comprehension processes. Decoding skill, however, requires special experiences that do not occur in the normal course of conversations. The characteristics of the writing systems may constitute constraints on the

Two experiments investigated the effects on auditory selection of varying distractor values in memory. Participants performed a set of control (single distractor) and distractor-variation (multiple distractors) tasks, classifying targets by pitch (Experiments 1A and 2) or loudness (Experiment 1B) while ignoring previously presented (and spatially separate) distractors. When both targets and distractors varied in pitch, the degree of variation among the distractors increasingly disrupted classification accuracy and reaction time to the targets. Physiologically, the degree of distractor variation boosted the N1 response to distractors, the P2 response to both targets and distractors, and the slow-wave response to targets (400-600 ms after stimulus onset). The results suggest that target representations are diminished in distinctiveness as distractors activate a wider range of the task-relevant continuum in working memory.

Behavioral studies have provided evidence that the processing of a musical target is faster and more accurate when it is harmonically related to the preceding prime context. We investigated the neural correlates of processing related and unrelated musical targets that were presented at the end of musical sequences. Participants were scanned with functional magnetic resonance imaging (fMRI) while performing speeded intonation judgments (consonant versus dissonant) on the target chords. Behavioral results acquired in the scanner replicated the facilitation effect of related over unrelated consonant targets. The overall activation pattern associated with target processing showed commonalities with networks previously described for target detection and novelty processing. The blood oxygen level-dependent (BOLD) signal linked to target processing revealed activation of bilateral inferior frontal regions (i.e. inferior frontal gyrus, frontal operculum, insula) that was stronger for unrelated than for related targets. We discuss our results with regard to the role of these areas in the processing and integration of temporal information.  (B. Tillmann).

We report a series a three psychophysical experiments designed to differentiate the contributions of the neocerebellar and prefrontal cortex to time perception. Comparison of patients with focal, unilateral neocerebellar or prefrontal lesions on temporal discrimination of Ž . 400-ms and 4-s intervals Expt. 1 indicated that neocerebellar damage impaired timing in both millisecond and seconds ranges, whereas prefrontal damage resulted in deficits that were robust only at the longer duration. Patients with prefrontal lesions, however, also Ž . exhibited working memory deficits on a non-temporal task Expt. 2 , biases in point of subjective equality indicative of attentional Ž . deficits, and were disproportionately sensitive to strategic manipulations in a long-duration discrimination task Expt. 3 . In contrast, the pervasive timing deficits of cerebellar patients were relatively insensitive to strategic support and could not be readily explained by general deficits in working memory or attention. These findings support the hypothesis that neocerebellar regions subserve a central timing mechanism, whereas the prefrontal cortex subserves supportive functions associated with the acquisition, maintenance, monitoring and organization of temporal representations in working memory. Such functions serve to bridge the output of the central timing mechanism with behavior. Together, these regions appear to participate in a working memory system involved in discrimination of durations extending from a few milliseconds to many seconds. q

In the stop-signal paradigm, fast responses are harder to inhibit than slow responses, so subjects must balance speed on the go task with successful stopping in the stop task. In theory, subjects achieve this balance by adjusting response thresholds for the go task, making proactive adjustments in response to instructions that indicate that relevant stop signals are likely to occur. We report five experiments that tested this theoretical claim, presenting cues that indicated whether or not stop signals were relevant for the next few trials. Subjects made proactive response-strategy adjustments in each experiment: diffusion-model fits showed that response threshold increased when subjects expected stop signals to occur, slowing go responses and increasing accuracy. Furthermore, our results show that subjects can make proactive responsestrategy adjustments on a trial-by-trial basis, suggesting a flexible cognitive system that can proactively adjust itself in changing environments.

Response properties of auditory cortical neurons measured in anesthetized preparations have provided important information on the physiological differences between neurons in different auditory cortical areas. Studies in the awake animal, however, have been much less common, and the physiological differences noted may reflect differences in the influence of anesthetics on neurons in different cortical areas. Because the behaving monkey is gaining popularity as an animal model in studies exploring auditory cortical function, it has become critical to physiologically define the response properties of auditory cortical neurons in this preparation. This study documents the response properties of single cortical neurons in the primary and surrounding auditory cortical fields in monkeys performing an auditory discrimination task. We found that neurons with the shortest latencies were located in the primary auditory cortex (AI). Neurons in the rostral field had the longest latencies and the narrowest intensity and frequency tuning, neurons in the caudomedial field had the broadest frequency tuning, and neurons in the lateral field had the most monotonic rate/level functions of the four cortical areas studied. These trends were revealed by comparing response properties across the population of studied neurons, but there was considerable variability between neurons for each response parameter other than characteristic frequency (CF) in each cortical area. Although the neuronal CFs showed a systematic spatial organization across AI, no such systematic organization was apparent for any other response property in AI or the adjacent cortical areas. The results of this study indicate that there are physiological differences between auditory cortical fields in the behaving monkey consistent with previous studies in the anesthetized animal and provide insights into the functional role of these cortical areas in processing acoustic information.

Psychophysiological studies with music have not examined what exactly in the music might be responsible for the observed physiological phenomena. The authors explored the relationships between 11 structural features of 16 musical excerpts and both self-reports of felt pleasantness and arousal and different physiological measures (respiration, skin conductance, heart rate). Overall, the relationships between musical features and experienced emotions corresponded well with those known between musical structure and perceived emotions. This suggests that the internal structure of the music played a primary role in the induction of the emotions in comparison to extramusical factors. Mode, harmonic complexity, and rhythmic articulation best differentiated between negative and positive valence, whereas tempo, accentuation, and rhythmic articulation best discriminated high arousal from low arousal. Tempo, accentuation, and rhythmic articulation were the features that most strongly correlated with physiological measures. Music that induced faster breathing and higher minute ventilation, skin conductance, and heart rate was fast, accentuated, and staccato. This finding corroborates the contention that rhythmic aspects are the major determinants of physiological responses to music.

With the growing prevalence of large databases of multimedia content, methods for facilitating rapid browsing of such databases or the results of a database search are becoming increasingly important. However, these methods are necessarily media dependent. We present a system for producing short, representative samples (or "audio thumbnails") of selections of popular music. The system searches for structural redundancy within a given song with the aim of identifying something like a chorus or refrain. To isolate a useful class of features for performing such structure-based pattern recognition, we present a development of the chromagram, a variation on traditional time-frequency distributions that seeks to represent the cyclic attribute of pitch perception, known as chroma. The pattern recognition system itself employs a quantized chromagram that represents the spectral energy at each of the 12 pitch classes. We evaluate the system on a database of popular music and score its performance against a set of "ideal" thumbnail locations. Overall performance is found to be quite good, with the majority of errors resulting from songs that do not meet our structural assumptions.

A broad consensus within the neuroscience of tinnitus holds that this audiologic condition is triggered by central deafferentation, mostly due to cochlear damage. The absence of audiometrically detectable hearing loss however poses a challenge to this rather generalizing assumption. The aim of this study was therefore to scrutinize cochlear functioning in a sample of tinnitus subjects audiometrically matched to a normal hearing control group. Two tests were applied: the Threshold Equalizing Noise (TEN) test and a pitch scaling task. To perform well on both tasks relatively normal functioning of inner hair cells is a requirement. In the TEN test the tinnitus group revealed a circumscribed increment of thresholds partially overlapping with the tinnitus spectrum. Abnormal slopes were observed in the pitch scaling task which indicated that tinnitus subjects, when presented with a high-frequency stimulus, relied heavily on input derived from lower-frequency inner hair cells (off-frequency listening). In total both results argue for the presence of a deafferentation also in tinnitus subjects with audiometrically normal thresholds and therefore favour the deafferentation assumption posed by most neuroscientific theories.

Using a habituation-recovery paradigm adapted to functional magnetic resonance imaging, we investigated the brain responses to syllables and tones in six right-handed male subjects. We opposed a standard condition (STD) in which the subjects were listening to homogeneous sequences of four identical stimuli, to a deviant condition (DEV) in which the fourth stimulus of the sequence differed in pitch or spectral content for tones and in the initial stop consonant for syllables. The corresponding runs alternated four rest periods with two STD and two DEV conditions. In addition to a marked rightward asymmetry in the primary and secondary auditory cortex for tones and a right inferior frontal activation for the tone condition where the deviant had increased spectral content, the experiment revealed differential activations in the left posterior superior temporal gyrus and in the left supramarginal gyrus. Activations within the left posterior superior temporal gyrus were observed for the DEV condition with tones and for the STD and DEV conditions with syllables. Activation within the inferior part of the left supramarginal gyrus was only observed for the DEV condition with syllables. The analysis of the decreases and increases in the BOLD signal across the STD, DEV, and rest conditions suggests that the left posterior superior temporal gyrus is implicated in the preattentive change detection of acoustic changes in speech as well as nonspeech stimuli, whereas the left supramarginal gyrus is more specifically engaged in the detection of changes in phonological units. 1999

The relative pitch of harmonic complex sounds, such as instrumental sounds, may be perceived by decoding either the fundamental pitch (f 0 ) or the spectral pitch (f SP ) of the stimuli. We classified a large cohort of 420 subjects including symphony orchestra musicians to be either f 0 or f SP 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 f 0 listeners, f SP 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.

Some scholars consider music to exemplify the classic criteria for a complex human adaptation, including universality, orderly development, and special-purpose cortical processes. The present account focuses on processing predispositions for music. The early appearance of receptive musical skills, well before they have obvious utility, is consistent with their proposed status as predispositions. Infants' processing of musical or music-like patterns is much like that of adults. In the early months of life, infants engage in relational processing of pitch and temporal patterns. They recognize a melody when its pitch level is shifted upward or downward, provided the relations between tones are preserved. They also recognize a tone sequence when the tempo is altered so long as the relative durations remain unchanged. Melodic contour seems to be the most salient feature of melodies for infant listeners. However, infants can detect interval changes when the component tones are related by small-integer frequency ratios. They also show enhanced processing for scales with unequal steps and for metric rhythms. Mothers sing regularly to infants, doing so in a distinctive manner marked by high pitch, slow tempo, and emotional expressiveness. The pitch and tempo of mothers' songs are unusually stable over extended periods. Infant listeners prefer the maternal singing style to the usual style of singing, and they are more attentive to maternal singing than to maternal speech. Maternal singing also has a moderating effect on infant arousal. The implications of these findings for the origins of music are discussed.

Background: Mismatch negativity (MMN) is an eventrelated brain potential that is sensitive to stimulus deviation from a repetitive pattern. The MMN is thought primarily to reflect the activity of sensory memory, with, at most, moderate influences of higher-level cognitive processes, such as attention. The MMN is reported to be reduced in patients with chronic schizophrenia. However, it is unknown whether MMN is reduced in patients with first-episode schizophrenia (at first hospitalization).

Communicating with one's environment requires efficient neural interaction between action and perception. Neural substrates ofsound perception and production are connected by the arcuate fasciculus (AF). While AF is known to be involved in language, its roles in non-linguistic functions are unexplored. Here we show that tone-deaf people, with impaired sound perception and production, have reduced AF connectivity. Diffusion tensor tractography and psychophysics were assessed in tone-deaf individuals and matched controls. Abnormally-reduced AF connectivity was observed in the tone-deaf. Furthermore, we observed relationships between AF and auditory-motor behavior: superior and inferior AF branches predict psychophysically-assessed pitch-discrimination and sound production-perception abilities respectively. This neural abnormality suggests that tone-deafness leads to a reduction in connectivity resulting in pitch-related impairments. Results support a dualstream anatomy of sound production and perception implicated in vocal communications. By identifying white-matter differences and their psychophysical correlates, results contribute to our understanding of how neural connectivity subserves behavior.

Adult subjects were presented with two auditory stimuli per trial, and their task was to decide which of the two was longer in duration. An adaptive psychophysical procedure was used. In Experiments 1, 2, and 4, the base duration was 50 msec, whereas in Experiment 3, the base duration was 1 sec. In Experiments 1, 2, and 4, it was found that filled intervals (continuous tones) were discriminated more accurately than empty intervals (with onset and offset marked by clicks). It was concluded that this difference was perceptual rather than cognitive in nature, since performance on filled and empty intervals was not affected by increasing cognitive load in a dual-task procedure (Experiment 2) but was affected by backward masking (Experiment 4). In contrast, the results of Experiment 3 showed that duration discriminationrf fIlled auditory intervals of longer duration was cognitively influenced, since performance was impaired by increasing cognitive load. Implications for notions of perceptual processing and timing mechanisms tanderlyingdifferences in duration discrimination with filled and empty intervals are discussed.

In adults, specific neural systems with right-hemispheric weighting are necessary to process pitch, melody, and harmony as well as structure and meaning emerging from musical sequences. It is not known to what extent the specialization of these systems results from longterm exposure to music or from neurobiological constraints. One way to address this question is to examine how these systems function at birth, when auditory experience is minimal. We used functional MRI to measure brain activity in 1-to 3-day-old newborns while they heard excerpts of Western tonal music and altered versions of the same excerpts. Altered versions either included changes of the tonal key or were permanently dissonant. Music evoked predominantly right-hemispheric activations in primary and higher order auditory cortex. During presentation of the altered excerpts, hemodynamic responses were significantly reduced in the right auditory cortex, and activations emerged in the left inferior frontal cortex and limbic structures. These results demonstrate that the infant brain shows a hemispheric specialization in processing music as early as the first postnatal hours. Results also indicate that the neural architecture underlying music processing in newborns is sensitive to changes in tonal key as well as to differences in consonance and dissonance. auditory cortex | functional MRI | neonates | emotion

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.

The familiar buzz of flying mosquitoes is an important mating signal, with the fundamental frequency of the female’s flight tone signaling her presence. In the yellow fever and dengue vector Aedes aegypti, both sexes interact acoustically by shifting their flight tones to match, resulting in a courtship duet. Matching is made not at the fundamental frequency of 400 hertz (female) or 600 hertz (male) but at a shared harmonic of 1200 hertz, which exceeds the previously known upper limit of hearing in mosquitoes. Physiological recordings from Johnston’s organ (the mosquito’s “ear”) reveal sensitivity up to 2000 hertz, consistent with our observed courtship behavior. These findings revise widely accepted limits of acoustic behavior in mosquitoes.

Purpose: We review a series of experiments aimed at studying pitch processing in music and speech. These studies were conducted with musician and non musician adults and children. We found that musical expertise improved pitch processing not only in music but also in speech. Demonstrating transfer of training between music and language has interesting applications for second language learning. We also addressed the issue of whether the positive effects of musical expertise are linked with specific predispositions for music or with extensive musical practice. Results of longitudinal studies argue for the later. Finally, we also examined pitch processing in dyslexic children and found that they had difficulties discriminating strong pitch changes that are easily discriminate by normal readers. These results argue for a strong link between basic auditory perception abilities and reading abilities. Methods: We used conjointly the behavioral method (Reaction Times and error rates) and the electrophysiological method (recording of the changes in brain electrical activity time-locked to stimulus presentation, Event-Related brain Potentials or ERPs). Results: A set of common processes may be responsible for pitch processing in music and in speech and these processes are shaped by musical practice. Conclusion: These data add evidence in favor of brain plasticity and open interesting perspectives for the remediation of dyslexia using musical training.

Visual stimuli are known to influence the perception of auditory stimuli in spatial tasks, giving rise to the ventriloquism effect. These influences can persist in the absence of visual input following a period of exposure to spatially disparate auditory and visual stimuli, a phenomenon termed the ventriloquism aftereffect. It has been speculated that the visual dominance over audition in spatial tasks is due to the superior spatial acuity of vision compared with audition. If that is the case, then the auditory system should dominate visual perception in a manner analogous to the ventriloquism effect and aftereffect if one uses a task in which the auditory system has superior acuity. To test this prediction, the interactions of visual and auditory stimuli were measured in a temporally based task in normal human subjects. The results show that the auditory system has a pronounced influence on visual temporal rate perception. This influence was independent of the spatial location, spe...

Musicians have a variety of perceptual and cortical specializations compared to non-musicians. Recent studies have shown that potentials evoked from primarily brainstem structures are enhanced in musicians, compared to non-musicians. Specifically, musicians have more robust representations of pitch periodicity and faster neural timing to sound onset when listening to sounds or both listening to and viewing a speaker. However, it is not known whether musician-related enhancements at the subcortical level are correlated with specializations in the cortex. Does musical training shape the auditory system in a coordinated manner or in disparate ways at cortical and subcortical levels? To answer this question, we recorded simultaneous brainstem and cortical evoked responses in musician and non-musician subjects. Brainstem response periodicity was related to early cortical response timing across all subjects, and this relationship was stronger in musicians. Peaks of the brainstem response evoked by sound onset and timbre cues were also related to cortical timing. Neurophysiological measures at both levels correlated with musical skill scores across all subjects. In addition, brainstem and cortical measures correlated with the age musicians began their training and the years of musical practice. Taken together, these data imply that neural representations of pitch, timing and timbre cues and cortical response timing are shaped in a coordinated manner, and indicate corticofugal modulation of subcortical afferent circuitry.

The neural basis of human pitch perception is not fully understood. It has been argued that the auditory cortices in the two hemispheres are specialized, such that certain right auditory cortical regions have a relatively finer resolution in the frequency domain than homologous regions in the left auditory cortex, but this concept has not been tested directly. Here, we used functional magnetic resonance imaging (fMRI) to test this specific prediction. Healthy volunteers were scanned while passively listening to pure-tone melodic-like sequences in which the pitch distance between consecutive tones was varied in a parametric fashion. As predicted, brain activation in a region of right lateral auditory cortex, corresponding to the planum temporale, was linearly responsive to increasing pitch distance, even across the fine changes in pitch. In contrast, the BOLD signal at the homologous left cortical region was relatively constant as a function of pitch distance, except at the largest pitch change. The results support the model of relative hemispheric specialization and indicate that the right secondary auditory cortex has a finer pitch resolution than the left.

We have previously shown that some visual motion areas can be specifically recruited by auditory motion processing in blindfolded sighted subjects . Specific activation of V5 brain area by auditory motion processing: an fMRI study. Brain Res. Cogn. Brain Res. 25, 650 -658]. The present fMRI study investigated whether auditory motion processing may recruit the same brain areas in early blind subjects. The task consisted of simultaneously determining both the nature of a sound stimulus (pure tone or complex sound) and the presence or absence of its movement. When a movement was present, blind subjects had to identify its direction. Auditory motion processing, as compared to static sound processing, activated the brain network of auditory and visual motion processing classically observed in sighted subjects. Accordingly, brain areas previously considered as specific to visual motion processing could be specifically recruited in blind people by motion stimuli presented through the auditory modality. This indicates that the occipital cortex of blind people could be organized in a modular way, as in sighted people. The similarity of these results with those we previously observed in sighted subjects suggests that occipital recruitment in blind people could be mediated by the same anatomical connections as in sighted subjects. D

This paper presents the design and the evaluation of a method to study prosodic features of discourse structure in unrestricted spontaneous speech. Past work has indicated that one of the major difficulties that discourse prosody analysts have to overcome is finding an independent specification of hierarchical discourse structure as to avoid circularity. Previous studies have tried to solve this problem by constraining the discourse or by basing segmentations on a specific discourse theory. The current investigation first explores the possibility of experimentally determining discourse boundaries in unrestricted speech. In a next stage, it is investigated to what extent boundaries obtained in this way correlate with specific prosodic variables: the features pause, pitch range, and type of boundary tone are studied as a function of discourse structure.

The afferent projections to the lateral superior olive (LSO) were examined with horseradish eroxidase, horseradish peroxidase-wheat germ agglutinin conjugate, l2 I-wheat germ agglutinin and tritiated leucine autoradiography, anterograde axonal degeneration, and 14C-2-deoxyglucose methods. The pathway to the ipsilateral LSO orginates in the spherical cells in anteroventral cochlear nucleus. Although some of the fibers pass above the lateral nucleus of the trapezoid body, most pass below it and turn at right angles to enter the LSO either directly through its ventral, lateral, or dorsal borders, or through its ventral or dorsal hilus. They end in unpolarized terminal fields throughout the LSO. Most if not all of these fibers are true collaterals of axons continuing across the midline in the trapezoid body. Verifying Held's (1893) finding of a major direct projection from the cochlear nucleus to the contralateral medial nucleus of the trapezoid body (MTB) and Rasmussen's ('46) finding of a major projection from the MTB to the LSO, the present results illustrate that this two-neuron pathway probably supplies all but a very small component of the relatively direct input to the LSO from the contralateral ear. This pathway originates in the globular cells of the ventral cochlear nucleus and relays mostly though not exclusively through the "principal cells" in the more rostra1 parts of the MTB. It terminates mostly in perisomal endings in unpolarized fields throughout the LSO, though most heavily within the (high frequency) medial and middle limbs and less heavily in the LSO's (low frequency) lateral limb. In addition to this indirect pathway, there is a small direct pathway to the contralateral LSO as suggested by Goldberg and Brown ('69) and Warr ('72, '82). This direct pathway to the contralateral LSO, like the direct ipsilateral pathway, probably originates in the spherical cell region of the ventral cochlear nucleus, crosses the midline in the trapezoid body, and terminates in a small circumscribed area within the LSO's ventromedial (high frequency) area. The 2-deoxyglucose method applied to cats in which the ipsilateral and contralateral pathways have been surgically isolated shows that each of the pathways converging on the LSO is topographically and tonotopically organized with the ipsilateral and the combined contralateral terminations in strict tonotopic register.