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Dissecting nonverbal auditory cortex asymmetry: an fMRI study.

The purpose of this study was to find a robust nonverbal paradigm to obtain reliable, reproducible auditory activation and characterize the nonverbal activation of the auditory cortex in regard to the Brodmann regions. The extent of localization and lateralization of activation was investigated utilizing functional magnetic resonance (fMR). Two tasks were used: monotonous repetitive stimuli of "double octaves" (DO) consisting of alternating four A with four C piano notes and a variated string of "sequential notes" (SN), which was a fast nonrepetitive sequence of piano notes. Eleven volunteers were investigated. The activation periods had a duration of 30 s, and presented every 30 s. All subjects demonstrated fMRI signal activity in the superior temporal gyrus (STG) involving the primary and secondary auditory cortex except one subject who showed no activation with the DO stimulus. SN elicited more activation than DO (p =<.03). The bulk activation for SN and DO was slightly greater in the right hemisphere, although the primary auditory area (Brodmann's 41) was better activated on the left p =<.001. Brodmann's area 22 was most frequently right-side dominant (p =.015, p =.017 for DO and SN, respectively). These findings appear to have implications in the examination of preverbal subjects.

Acoustic Stimulation↗

A comparison of neuron response properties in areas A1 and CM of the marmoset monkey auditory cortex: tones and broadband noise.

The purpose of this study was to compare response properties of two adjacent areas of the marmoset monkey auditory cortex. Multiunit responses to 50 ms tones and broadband noise bursts (BBN) were recorded in the core area, A1, and the caudomedial belt area, CM, of ketamine-anesthetized animals. Neurons in A1 and CM exhibited robust low-threshold short-latency responses to BBN and tones, whereas neurons in adjoining lateral belt areas were poorly responsive or unresponsive to tones and noise. Except for a population of broadly tuned units in CM, the characteristic frequency (CF) could be determined for all recording sites in A1 and CM. Both areas were tonotopically organized and shared a high CF border. Whereas the tonotopic gradient in A1 was smooth and continuous across the field, the gradient in CM was discontinuous, and the intermediate CF range was underrepresented. For BBN stimuli, rate level functions were largely monotonic in A1 and CM. Response profiles were also similar in both areas. As a population, neurons in CM were distinguished from A1 by significantly shorter response latencies, lower thresholds, and broader tuning bandwidth at higher intensities. The results indicated that, while A1 and CM represent anatomically and physiologically distinct areas, their response profiles under anesthesia overlapped considerably compared with the lateral belt areas. Therefore refinements of current models of the primate auditory cortex may be needed to account for differences in organization among the auditory belt areas.

Acoustic Stimulation↗

Responses of the human auditory cortex to vowel onset after fricative consonants.

Neuromagnetic responses to different auditory stimuli (noise bursts and short speech stimuli) were mapped over both hemispheres of seven healthy subjects. The results indicate that a particular acoustic feature of speech, vowel onset after voiceless fricative consonants, evokes a prominent response in the human supratemporal auditory cortex. Although the observed response seems to be specific to acoustic rather than phonetic characteristics of the stimuli, it might reflect feature detection essential for further speech processing.

Auditory Cortex↗

Encoding of learned importance of sound by magnitude of representational area in primary auditory cortex.

We hypothesized that learning-induced representational expansion in the primary auditory cortex (AI) directly encodes the degree of behavioral importance of a sound. Rats trained on an operant auditory conditioning task were variably motivated to the conditioned stimulus (CS) through different levels of water deprivation. Mean performance values correlated with deprivation level, validating them as a measure of the overall control and, therefore, behavioral importance of the CS. Electrophysiological mapping revealed expanded representations of the CS, compared with other frequencies in experimental subjects, but not in naive or visually trained controls that received noncontingent CS tones. Importantly, representational area showed a significant positive correlation with mean performance levels for only the CS band, with significant effects for relative area in contrast to only modest changes in absolute area. CS representational expansion was asymmetric into high-frequency zones, thus performance level also was significantly correlated with the relative anterior-posterior location of the enlarged representation. An increased representation of low frequencies, related to the acoustic spectrum of the reward delivery equipment, also was discovered in both experimental and control trained subjects, supporting the conclusion that behaviorally important sounds gain representational area. Furthermore, there was a surprising reduction in total AI area for the experimental and control groups, compared with untrained naive subjects, indicating that the functional dimensions of AI are not fixed. Overall, the findings support the encoding of acquired stimulus importance based on representational size in AI.

Acoustic Stimulation↗

Sensory-motor interaction in the primate auditory cortex during self-initiated vocalizations.

Little is known about sensory-motor interaction in the auditory cortex of primates at the level of single neurons and its role in supporting vocal communication. The present study investigated single-unit activities in the auditory cortex of a vocal primate, the common marmoset (Callithrix jacchus), during self-initiated vocalizations. We found that 1) self-initiated vocalizations resulted in suppression of neural discharges in a majority of auditory cortical neurons. The vocalization-induced inhibition suppressed both spontaneous and stimulus-driven discharges. Suppressed units responded poorly to external acoustic stimuli during vocalization. 2) Vocalization-induced suppression began several hundred milliseconds prior to the onset of vocalization. 3) The suppression of cortical discharges reduced neural firings to below the rates expected from a unit's rate-level function, adjusted for known subcortical attenuation, and therefore was likely not entirely caused by subcortical attenuation mechanisms. 4) A smaller population of auditory cortical neurons showed increased discharges during self-initiated vocalizations. This vocalization-related excitation began after the onset of vocalization and is likely the result of acoustic feedback. Units showing this excitation responded nearly normally to external stimuli during vocalization. Based on these findings, we propose that the suppression of auditory cortical neurons, possibly originating from cortical vocal production centers, acts to increase the dynamic range of cortical responses to vocalization feedback for self monitoring. The excitatory responses, on the other hand, likely play a role in maintaining hearing sensitivity to the external acoustic environment during vocalization.

Acoustic Stimulation↗

Phonetic invariance in the human auditory cortex.

Neuromagnetic signals evoked by synthesized syllables (/bae/ and /gae/) were recorded over the left auditory cortex of healthy humans. The fundamental frequencies of the syllables varied as if the same speaker had pronounced them at 16 different pitches. Specific mismatch responses to infrequent syllables among frequent syllables of the other type indicated that phonetically invariant information had been extracted at the level of the auditory cortex from the extensive irrelevant pitch variation. Such a detection mechanism is necessary for perceiving speech sounds in natural situations with a great deal of acoustic variation present.

Acoustic Stimulation↗

Word length modulates neural activity in auditory cortex during covert object naming.

Several lines of evidence show that posterior portions of left auditory cortex participate in aspects of speech production. A current hypothesis is that these regions play a specific role in processing phonological codes. We used event-related fMRI to test this hypothesis. Subjects covertly named objects that had names varying in length from one to four syllables. Behavioral data, both in previous work and in the present study, show an increase in naming reaction time as words get longer, reflecting the increase in phonological load. If left posterior auditory areas participate in phonological aspects of production, word length should modulate neural activity in this region. Object naming activated the left planum temporale in each subject, confirming previous findings of auditory cortex involvement in speech production. The timecourse of activation in this region showed a length effect, consistent with its hypothesized role in phonological processing. Similar effects were also observed in premotor cortex, but not in occipital-temporal cortex, which presumably supports visual processing of the pictures to be named.

Auditory Cortex↗

Plastic changes in ipsi-contralateral differences of auditory cortex and inferior colliculus evoked potentials after injury to one ear in the adult guinea pig.

In normal adult guinea pigs, evoked potentials recorded at the ipsilateral auditory cortex to monaural high-frequency acoustic stimuli present higher thresholds and lower amplitudes than at the contralateral cortex; in the inferior colliculus, such ipsi-contralateral differences (ICDs) are smaller than in the auditory cortex. Changes in the ICDs were studied after opposite ear injury. Following quasi-complete hair cell destruction induced by sisomicin injection into the contralateral inner ear, threshold ICDs almost disappeared after about two to six days and ipsilateral amplitudes progressively increased in two to three weeks. The occurrence of ICDs at higher auditory centers revealed in this study, indicates peculiar processing of high frequency stimuli in normal guinea pigs. The alteration of ICDs after opposite ear impairment provides a new possibility to study the auditory plasticity in adult animals.

Acoustic Stimulation↗

Dynamic movement of N100m dipoles in evoked magnetic field reflects sequential activation of isofrequency bands in human auditory cortex.

OBJECTIVE: To investigate spatiotemporal features of the isofrequency bands for 400 and 4000 Hz tones in human auditory cortex and on the hemispheric differences in the arrangement of isofrequency bands. METHODS: We recorded auditory evoked magnetic fields (AEFs) to 400 or 4000 Hz tone pips presented at right or left ear from 31 normal subjects. The dipole location for the N100m sources was successively calculated from the AEFs obtained from the hemisphere contralateral to the stimulated ear. RESULTS: In the right hemisphere, the current sources for 400 and 4000 Hz moved toward the anterolateral direction before the N100m peak, showing parallel arrangement of the isofrequency bands (4000 Hz in medial location). In the left hemisphere, the movement direction of 400 Hz dipoles was anterolateral, while that of 4000 Hz dipoles was lateral. CONCLUSIONS: This difference in the organization of isofrequency bands between right and left auditory cortices reflects distinct functional roles in auditory information processing such as pitch vs. language discrimination. SIGNIFICANCE: This work is the first to disclose isofrequency bands in human auditory cortex based on the analysis of magnetoencephalography.

Acoustic Stimulation↗

Kindling limits the interictal neuronal temporal response properties in cat primary auditory cortex.

PURPOSE: The present study examined the effect of electrical kindling on the interictal temporal response properties of single units recorded from primary auditory cortex (AI) of the adult cat. METHODS: Cats were permanently implanted with electrodes in AI, kindled twice daily for 40 sessions, and the contralateral AI was subsequently mapped. Kindling stimulation consisted of 1-s trains of biphasic square-wave pulses applied at a frequency of 60 Hz, 100 microA above the afterdischarge (AD) threshold. The EEG activity was recorded during each kindling session, and the behavioral manifestation was scored. Subsequent to kindling, multiple single-unit responses were recorded under ketamine anesthesia in response to 1-s-long periodic click trains, with click rates between 2 and 64 Hz. Neuronal responses were characterized according to their ability to respond in time-locked fashion to the clicks. RESULTS: Kindling stimulation resulted in progression of the AD characteristics and seizure behavior, with six of 10 kindled cats reaching a fully generalized state. In the fully kindled cats, the best modulation frequencies and limiting following rates for the single-unit responses were significantly lower compared with those of naive and sham controls. CONCLUSIONS: Repeated epileptiform activity interferes with temporal processing in cat auditory cortex in the interictal state. This may have implications for people with epileptic foci in auditory-related areas.

Acoustic Stimulation↗

Integration of touch and sound in auditory cortex.

To form a coherent percept of the environment, our brain combines information from different senses. Such multisensory integration occurs in higher association cortices; but supposedly, it also occurs in early sensory areas. Confirming the latter hypothesis, we unequivocally demonstrate supra-additive integration of touch and sound stimulation at the second stage of the auditory cortex. Using high-resolution fMRI of the macaque monkey, we quantified the integration of auditory broad-band noise and tactile stimulation of hand and foot in anaesthetized animals. Integration was found posterior to and along the lateral side of the primary auditory cortex in the caudal auditory belt. Integration was stronger for temporally coincident stimuli and obeyed the principle of inverse effectiveness: greater enhancement for less effective stimuli. These findings demonstrates that multisensory integration occurs early and close to primary sensory areas and--because it occurs in anaesthetized animals--suggests that this integration is mediated by preattentive bottom-up mechanisms.

Animals↗

Long lasting attenuation by prior sounds in auditory cortex of awake primates.

How the brain responds to sequences of sounds is a question of great relevance to a variety of auditory perceptual phenomena. We investigated how long the responses of neurons in the primary auditory cortex of awake monkeys are influenced by the previous sound. We found that responses to the second sound of a two-sound sequence were generally attenuated compared to the response that sound evoked when it was presented first. The attenuation remained evident at the population level even out to inter-stimulus intervals (ISIs) of 5 s, although it was of modest size for ISIs >2 s. Behavioral context (performance versus non-performance of a visual fixation task during sound presentation) did not influence the results. The long time course of the first sound's influence suggests that, under natural conditions, neural responses in auditory cortex are rarely governed solely by the current sound.

Acoustic Stimulation↗

Tone responses in core versus belt auditory cortex in the developing chinchilla.

Single-unit responses to tone pip stimuli were isolated from numerous microelectrode penetrations of core primary auditory cortex (AI) and a dorsocaudal (DC) belt region in the ketamine-anesthetized chinchilla (laniger). Results are reported at postnatal day 3 (P3), P15, P30, and from adult animals. The AI core could be distinguished from the DC belt on the basis of its strict tonotopic organization, evident in all chinchillas studied (including the youngest). Averaged by age group and compared to their core counterparts, belt neurons generally had similar absolute (spike rate) thresholds and onset latencies (at a given sound pressure level), but lower maximum spike rates, broader tuning bandwidths, and more complex (multipeaked) receptive fields. Most notably, the fraction of complex belt units in the near-newborn (P3) group was high (approximately 50%), and did not systematically increase with age, while that of complex core units was approximately 10% at P3 and increased steadily to about 40% in adulthood. These results provide further evidence to support the hypothesis that, at least to some extent, core and belt auditory cortex may constitute parallel processing streams which represent different aspects of complex acoustic stimuli.

Aging↗

Projections from auditory cortex contact cells in the cochlear nucleus that project to the inferior colliculus.

Anterograde and retrograde tracing techniques were combined to determine whether auditory cortical axons contact cells in the cochlear nucleus that project to the inferior colliculus. FluoroRuby or fluorescein dextran was injected into auditory cortex to label cortical axons by anterograde transport. Different fluorescent tracers (Fast Blue, FluoroGold, FluoroRuby or fluorescein dextran) were injected into one or both inferior colliculi to label cells in the cochlear nucleus. After 12-15 days, the brain was processed for fluorescence microscopy and the cochlear nuclei were examined for apparent contacts between cortical axons and retrogradely labeled cochlear nucleus cells. The results suggest that axons from the ipsilateral or contralateral cortex contact fusiform and giant cells in the dorsal cochlear nucleus and multipolar cells in the ventral cochlear nucleus that project directly to the inferior colliculus. The contacts occur on cell bodies and dendrites. The target cells in the cochlear nucleus include cells that project ipsilaterally, contralaterally or bilaterally to the inferior colliculus. The results suggest that auditory cortex is in a position to exert direct effects on the monaural pathways that ascend from the cochlear nucleus.

Animals↗

Functional MR spectroscopy of the auditory cortex in healthy subjects and patients with sudden hearing loss.

PURPOSE: To use MR spectroscopy to study the biochemical changes produced by auditory stimuli in patients with sudden sensorineural hearing loss and to compare these findings with the biochemical changes seen in healthy volunteers. METHODS: Single-voxel MR spectroscopy was used to study biochemical changes in the auditory cortex in 11 control subjects and 19 patients with sudden sensorineural hearing loss. MR spectroscopic signals were measured during three different sound conditions (scanner noise, music, and sirens). RESULTS: A lower MR spectroscopic lactate signal was observed in control subjects during the music stimulus than during the other sound conditions. This music-induced lactate change was not observed in patients with hearing loss. The other proton metabolites (choline, creatine, N-acetylaspartate [NAA]) remained stable during the different auditory stimuli. However, the NAA/creatine ratio was higher in the auditory cortex of patients than in the control subjects, and was not dependent on the sound condition. CONCLUSION: The detection of stimulus-induced and stable biochemical MR spectroscopic changes in patients with hearing loss may be useful in assessing disease activity.

Acoustic Stimulation↗

Optical imaging of intrinsic signals in chinchilla auditory cortex.

We have assessed sound frequency and intensity responses in primary auditory cortex of the (ketamine) anesthetized chinchilla using optical imaging of intrinsic signals. Temporal cortex was exposed via a 10-mm craniotomy and a windowed chamber was mounted. A 4-second period of gated tones (10 ms rise/fall; 50 ms plateau; 10/s) was presented to the contralateral ear at levels between 0 and 80 dB SPL. The cortical surface was illuminated with 540 nm light and video images captured in 0.5-second bins for 7.5 s (Imager 2001; Optical Imaging). Intrinsic signals were first apparent 0.5-1 s after stimulus onset, and were maximal after 3-4 s; they decayed over several seconds. The cortical area in which intrinsic activity was detected corresponded closely with electrophysiologically defined AI cortex. Intrinsic signals can reliably be detected to stimuli at 30-40 dB SPL, and in general, the area of intrinsic signal activity tends to expand with increasing stimulation level. Using stimulation levels of 80 dB SPL, we show that low-frequency stimuli (0.5-1 kHz) evoke intrinsic signals in anterior areas whilst posterior areas are activated by high-frequency stimuli (e.g. 16 kHz). Thus a low- to high-frequency tonotopic organization is seen along this axis.

Animals↗

Acquired word deafness, and the temporal grain of sound representation in the primary auditory cortex.

This paper explores the nature of the processing disorder which underlies the speech discrimination deficit in the syndrome of acquired word deafness following from pathology to the primary auditory cortex. A critical examination of the evidence on this disorder revealed the following. First, the most profound forms of the condition are expressed not only in an isolation of the cerebral linguistic processor from auditory input, but in a failure of even the perceptual elaboration of the relevant sounds. Second, in agreement with earlier studies, we conclude that the perceptual dimension disturbed in word deafness is a temporal one. We argue, however, that it is not a generalized disorder of auditory temporal processing, but one which is largely restricted to the processing of sounds with temporal content in the milliseconds to tens-of-milliseconds time frame. The perceptual elaboration of sounds with temporal content outside that range, in either direction, may survive the disorder. Third, we present neurophysiological evidence that the primary auditory cortex has a special role in the representation of auditory events in that time frame, but not in the representation of auditory events with temporal grains outside that range.

Animals↗

[Functional imaging of the auditory cortex: role of magnetoencephalography].

We present the different methods for detecting cerebral activity in the auditory cortex. Positron emission tomography (PET) and functional MRI (fMRI) measure such activity indirectly by calculating the blood flow rate or the consumption of oxygen. The direct methods of detection record the electrical or magnetic activity by eletroencephalography (EEG), or magnetoencephalography (MEG), respectively. The aim of this study was to define the role of MEG amongst these different techniques using the data from recordings of evoked magnetic fields in 5 healthy subjects. The localizations demonstrated the tonotopic organization of the auditory cortex, with high-pitched sounds showing a more medial cortical projection than low-pitched sounds. These various techniques are complementary. PET allows a pharmacological study of the cortex, and could be used in patients with cochlear implants. FMRI is non-invasive, and has a high spatial resolution. EEG has an excellent temporal resolution, and EEG recordings do not require major equipment or infrastructure. MEG has a better spatial resolution, with the same temporal resolution, as EEC. MEG is particularly useful in the localization of the cortical generators of middle-latency auditory evoked responses.

Adult↗