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Event-related magnetic fields in the auditory cortex of man during unilateral movements: a discriminant function analysis.

It is often assumed that sensorimotor coordination is a feature of the sensorimotor areas of the neocortex only. The purpose of the present study was to examine how this phenomenon is reflected in the auditory cortex of man. Ten subjects were engaged in a stimulus-reaction paradigm, in which each of two acoustical tones was associated to either of two motor reactions. Magnetic fields recorded with a 122-channel magnetometer were modelled by current dipoles. The spatial coordinates as well as the amplitudes of the dipoles were analyzed from 90 to 110 ms after stimulus onset using discriminant analysis. The results suggest that the dipole trajectory in the auditory cortex of the right hemisphere and amplitudes of the dipoles in the auditory cortex of the left hemisphere already 90-110 ms after the beginning of the stimulus could be affected not only by physical features of the stimulus, but also by the motor task required as a reaction.

Acoustic Stimulation↗

Neural correlates of gap detection and auditory fusion in cat auditory cortex.

Responses were recorded from 130 single neurones in the primary auditory cortex of 12 ketamine-anaesthetized cats in response to double-click stimuli, to a /ba/-/pa/ phoneme continuum and to gaps inserted early (after 5 ms) and late (after 500 ms) in a 1 s duration noiseburst. Stimulus levels were between 45 and 75 dB SPL. Neural detection threshold for the 'late gap' was less than 5 ms. For the double click and 'early gap' stimuli thresholds were between 40 and 50 ms, whereas the phoneme continuum threshold for voice-onset-time (VOT) was between 10 and 25 ms. The 'late gap' and VOT thresholds are similar to psychophysical gap detection and the /ba/-/pa/ categorical perception boundary respectively.

Acoustic Stimulation↗

Neuroplasticity of auditory cortex after stape surgery for otosclerosis: a magnetoencephalographic study.

Aim of the present study was to investigate the tonotopic reorganization of the primary auditory cortex in otosclerotic patients following functional stapedioplasty. Characteristics of auditory cortex activation have been evaluated in a series of 10 otosclerotic patients before and after surgery. In these patients, a magnetoencephalographic recording of evoked magnetic fields has been performed by means of tone-burst monoaural stimulation with frequency octaves between 250 and 2000 Hz. Brain topography of cortex response generators (wave N100m) in patients with otosclerosis has been compared with that observed in a control group of 10 healthy subjects: changes before and after surgery have also been correlated with the functional result as far as concerns improvement in hearing. A significant reduction has been observed in the cortical tonotopic extension in response to the acoustic stimulus in patients "pre-surgery" in comparison with controls: after surgery, tonotopic mapping showed an increase, dimensions becoming comparable to those in control subjects. This increase in size was found to be significantly correlated with duration of the post-operative period. Data emerging from the present study suggest that the cortical auditory areas in man are involved in a "plastic" functional reorganization following changes in the receptor or peripheral deprivation. Reduction in the cortical tonotopic mapping resulting from prolonged lowering of auditory "input" is modified by reorganization of the cortex after the recovery of auditory function: this process occurs over a period of a few weeks.

Adult↗

Consonance and dissonance of musical chords: neural correlates in auditory cortex of monkeys and humans.

Some musical chords sound pleasant, or consonant, while others sound unpleasant, or dissonant. Helmholtz's psychoacoustic theory of consonance and dissonance attributes the perception of dissonance to the sensation of "beats" and "roughness" caused by interactions in the auditory periphery between adjacent partials of complex tones comprising a musical chord. Conversely, consonance is characterized by the relative absence of beats and roughness. Physiological studies in monkeys suggest that roughness may be represented in primary auditory cortex (A1) by oscillatory neuronal ensemble responses phase-locked to the amplitude-modulated temporal envelope of complex sounds. However, it remains unknown whether phase-locked responses also underlie the representation of dissonance in auditory cortex. In the present study, responses evoked by musical chords with varying degrees of consonance and dissonance were recorded in A1 of awake macaques and evaluated using auditory-evoked potential (AEP), multiunit activity (MUA), and current-source density (CSD) techniques. In parallel studies, intracranial AEPs evoked by the same musical chords were recorded directly from the auditory cortex of two human subjects undergoing surgical evaluation for medically intractable epilepsy. Chords were composed of two simultaneous harmonic complex tones. The magnitude of oscillatory phase-locked activity in A1 of the monkey correlates with the perceived dissonance of the musical chords. Responses evoked by dissonant chords, such as minor and major seconds, display oscillations phase-locked to the predicted difference frequencies, whereas responses evoked by consonant chords, such as octaves and perfect fifths, display little or no phase-locked activity. AEPs recorded in Heschl's gyrus display strikingly similar oscillatory patterns to those observed in monkey A1, with dissonant chords eliciting greater phase-locked activity than consonant chords. In contrast to recordings in Heschl's gyrus, AEPs recorded in the planum temporale do not display significant phase-locked activity, suggesting functional differentiation of auditory cortical regions in humans. These findings support the relevance of synchronous phase-locked neural ensemble activity in A1 for the physiological representation of sensory dissonance in humans and highlight the merits of complementary monkey/human studies in the investigation of neural substrates underlying auditory perception.

Acoustic Stimulation↗

Neuronal connections in the primary auditory cortex: an electrophysiological study in the cat.

Neuronal connections in the primary auditory cortex (AI) of the cat were studied electrophysiologically by using intracellular recording techniques. Fast-conducting fibers from the medial geniculate nucleus (MG) projected monosynaptically onto AI neurons in layers III-VI (mainly in layer IV), whereas slow-conducting MG-fibers projected monosynaptically onto AI neurons in layer I. AI neurons which received monosynaptic inputs from the auditory association cortices (AII and Ep) and/or from the contralateral AI were distributed in all layers of the AI; the commissural fibers from the contralateral AI were divided into fast- and slow-conducting ones. AI neurons were categorized into seven types: type I neurons which received monosynaptic inputs from slow-conducting MG-fibers were located in layer I. Type II neurons which received polysynaptic inputs from the MG were located in layers II-VI. Type III neurons which sent their axons to the AII or Ep were mainly located in layer III. Type IV neurons which sent their axons to the contralateral AI were located mainly in layer III. Type V neurons which received monosynaptic inputs from fast-conducting MG-fibers were located mainly in layer IV. Type VI neurons which projected onto the inferior colliculus were located in the upper part of the layer V. Type VII neurons which projected onto the MG were located in layers V and VI.

Animals↗

Development of reorganization of the auditory cortex caused by fear conditioning: effect of atropine.

Reorganization of the frequency map in the central auditory system is based on shifts in the best frequencies (BFs; hereafter, BF shifts), together with the frequency-response curves, of auditory neurons. In the big brown bat, conditioning with acoustic stimulation followed by electric leg-stimulation causes BF shifts of collicular and cortical neurons. The collicular BF shift develops quickly and is short term, whereas the cortical BF shift develops slowly and is long term. The acetycholine level in the auditory cortex must be high during conditioning to develop these BF shifts. We studied the effect of atropine (an antagonist of muscarinic acetylcholine receptors) applied to the auditory cortex on the development of the long-term cortical BF shift in the awake bat caused by a 30-min conditioning session. We found 1) the cortical BF shift starts to develop approximately 15 min after the onset of the conditioning, gradually increases over 60 min, and reaches a plateau, 2) the cortical BF shift changes from short to long term approximately 45 min after the onset of the conditioning, 3) the cortical BF shift can plateau at different frequencies between the BF of a given neuron in the control condition and the frequency of the conditioning tone, 4) the maximum BF shift is determined approximately 70 min after the onset of the conditioning, and 5) acetylcholine plays an important role in the development of the cortical BF shift. Its role ends approximately 180 min after the onset of the conditioning.

Acoustic Stimulation↗

Binaural interaction revisited in the cat primary auditory cortex.

The binaural interactions of neurons were studied in the primary auditory cortex (AI) of barbiturate-anesthetized cats with a matrix of binaural tonal stimuli varying in both interaural level differences (ILD) and average binaural level (ABL). The purpose of this study was to determine: 1) the distribution of preferred binaural combinations (PBCs) of a large population of neurons and its relationships with binaural interactions and binaural monotonicity; 2) whether monaural responses are predictive of binaural responses; and 3) whether there is a restricted set of representative binaural stimulus configurations that could effectively classify the binaural interactions. Binaural interactions were often diverse in the matrix and dependent on both ABL and ILD. Compared with previous studies, a higher proportion of mixed binaural interaction type and a lower proportion of EO/I type were found. No monaural neurons were found. Binaural responses often differed from monaural responses in the number of spikes and/or the form of the response functions. The PBCs of the majority of EO and PB neurons were in the contralateral field and midline, respectively. However, the PBCs of EE units were evenly distributed across the contralateral and ipsilateral fields. The majority of the nonmonotonic neurons responded most strongly to lower ABLs, whereas the majority of monotonic neurons responded most strongly to higher ABLs. This study demonstrated that in AI a restricted set of binaural stimulus configurations is not sufficient to reveal the binaural responses properties. Also, monaural responses are not predictive of binaural responses.

Acoustic Stimulation↗

Factors shaping the tone level sensitivity of single neurons in posterior field of cat auditory cortex.

1. The posterior field (field P) of the cat's auditory cortex contains a higher proportion of neurons whose response/level functions for characteristic frequency (CF) tones are nonmonotonic than does the primary field (AI). The general purpose of the present study is to assess whether the response/level functions of field P neurons are generated by the same mechanisms as those of cells in AI. All of the data came from single neurons in the cortices of barbiturate-anesthetized cats, to which we presented tonal stimuli through sealed, calibrated stimulating systems. 2. We obtained quantitative data from 123 neurons, of which 108 were located in field P. Of the 108 field P cells, 70% had nonmonotonic response/level functions for 5-ms rise time tones of CF. For cells of any given CF, both CF thresholds and best SPLs (i.e., SPLs associated with maximal responses) varied widely. A correlation analysis revealed that a linear relation between best SPL and CF threshold accounted for 73% of the data variance in the association between those response variables. An analysis of data from 83 nonmonotonic cells in AI revealed a similar relation. 3. Field P neurons whose response/level functions were non-monotonic for 5-ms rise time CF tones became even more narrowly tuned to SPL when the rise time of the tone bursts was reduced to 1 ms. Lengthening the rise time to 20 ms reduced or eliminated the SPL tuning in almost all of these neurons. The general form of monotonic tone response/level functions was commonly unaffected by variation in signal rise time. In a few instances, cells with monotonic response/level functions for 5- and 20-ms rise time tones developed nonmonotonic functions for 1-ms rise time tones. 4. Field P neurons with nonmonotonic response/level functions for CF tones usually failed to respond to wideband noise pulses, or, less commonly, responded to noise only at low SPLs. In contrast, field P cells with a monotonic response to CF tones usually responded monotonically to noise. 5. The minimal mean first-spike latencies of field P neurons were generally longer than those of AI cells studied under similar conditions. The precision of first-spike timing, measured using the SD of the mean first-spike latency, was commonly poorer than that of AI cells. 6. The properties of field P cells followed the same rules as those seen in AI.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

Speaking modifies voice-evoked activity in the human auditory cortex.

The voice we most often hear is our own, and proper interaction between speaking and hearing is essential for both acquisition and performance of spoken language. Disturbed audiovocal interactions have been implicated in aphasia, stuttering, and schizophrenic voice hallucinations, but paradigms for a noninvasive assessment of auditory self-monitoring of speaking and its possible dysfunctions are rare. Using magnetoencephalograpy we show here that self-uttered syllables transiently activate the speaker's auditory cortex around 100 ms after voice onset. These phasic responses were delayed by 11 ms in the speech-dominant left hemisphere relative to the right, whereas during listening to a replay of the same utterances the response latencies were symmetric. Moreover, the auditory cortices did not react to rare vowel changes interspersed randomly within a series of repetitively spoken vowels, in contrast to regular change-related responses evoked 100-200 ms after replayed rare vowels. Thus, speaking primes the human auditory cortex at a millisecond time scale, dampening and delaying reactions to self-produced "expected" sounds, more prominently in the speech-dominant hemisphere. Such motor-to-sensory priming of early auditory cortex responses during voicing constitutes one element of speech self-monitoring that could be compromised in central speech disorders.

Adult↗

Functional organization of auditory cortex in the mongolian gerbil (Meriones unguiculatus). II. Tonotopic 2-deoxyglucose.

The tonotopic organization of the auditory cortex in the Mongolian gerbil was mapped with 2-deoxyfluoro-D-glucose (2DG) using narrow-band frequency-modulated tones of different centre frequency (FM tones) and tones periodically alternating between two different frequencies (alternating tones) as stimuli. Continuous tone bursts of a constant frequency and repetition rate were used in initial experiments. Continuous tones produced 2DG patterns similar to those observed in animals that were not specifically stimulated. With tone bursts of constant frequency and repetition rate variable patterns were observed, some of which could be interpreted only in retrospect in the light of results obtained with FM tones and alternating tones. These stimuli, in contrast, produced differential metabolic responses which in conjunction with 2DG data from monaural animals and electrophysiological data made it possible to distinguish a primary auditory field AI with its dorsal region Ald, an anterior auditory field AAF, a ventral field V, a dorsoposterior field DP and a ventroposterior field VP, a dorsal field D, and in addition an anteroventral field AV. In the largest field (AI) and the smaller rostrally adjacent field AAF, frequency-specific dorsoventral bands of labelling (isofrequency contours) were mapped quantitatively. Bands shifted as a function of frequency relative to each other and to an independent spatial reference line in the lateral hippocampus. Spatial analysis of the single bands obtained with FM tones, and of the double bands obtained with alternating tones in both fields, revealed roughly mirror-imaged tonotopic maps of AI and AAF. In AI the progression from low to high frequencies was from caudal to rostral and in AAF the gradient was reversed, leading to a common high-frequency border of the two fields. In AI, the spatial resolution for frequencies below 16 kHz was in similar intervals per octave and higher for frequencies below 1 kHz. AI showed a somewhat higher spatial resolution for frequencies (at least below 1 kHz) as well as longer isofrequency contours than AAF. The 2-deoxyglucose patterns provided average tonotopic maps and topological data on various fields, as well as reliable landmarks in the gerbil's auditory cortex.

Acoustic Stimulation↗

Habituation produces frequency-specific plasticity of receptive fields in the auditory cortex.

Associative learning produces conditioned stimulus (CS)-specific plasticity of frequency receptive fields (RFs) in the auditory cortex; responses to the CS frequency are increased, whereas responses to other frequencies are decreased. This study determined the effects of habituation on the RF of neurons in the auditory cortex of the guinea pig (Cavia porcellus). One frequency was presented repeatedly (REP) followed by redetermination of the RF. After REP, 26/36 (72%) RFs exhibited a substantial reduction (70-75%) of response to the repeated frequency, and this was highly specific (bandwidth less than 0.125 octave). This RF plasticity involves an initial decrease in response during REP but does not require attenuated responses at the end of REP. Incubation (i.e., development over time after cessation of REP) and long-term frequency-specific effects are evident. Thus, habituation induces a specific change in the processing of frequency information rather than a general reduction in responsivity.

Acoustic Stimulation↗

Auditory cortex of the rufous horseshoe bat: 1. Physiological response properties to acoustic stimuli and vocalizations and the topographical distribution of neurons.

The extent and functional subdivisions of the auditory cortex in the echolocating horseshoe bat, Rhinolophus rouxi, were neurophysiologically investigated and compared to neuroarchitectural boundaries and projection fields from connectional investigations. The primary auditory field shows clear tonotopic organization with best frequencies increasing in the caudorostral direction. The frequencies near the bat's resting frequency are largely over-represented, occupying six to 12 times more neural space per kHz than in the lower frequency range. Adjacent to the rostral high-frequency portion of the primary cortical field, a second tonotopically organized field extends dorsally with decreasing best frequencies. Because of the reversed tonotopic gradient and the consistent responses of the neurons, the field is comparable to the anterior auditory field in other mammals. A third tonotopic trend for medium and low best frequencies is found dorsal to the caudal primary field. This area is considered to correspond to the dorsoposterior field in other mammals. Cortical neurons had different response properties and often preferences for distinct stimulus types. Narrowly tuned neurons (Q10dB > 20) were found in the rostral portion of the primary field, the anterior auditory field and in the posterior dorsal field. Neurons with double-peaked tuning curves were absent in the primary area, but occurred throughout the dorsal fields. Vocalization elicited most effectively neurons in the anterior auditory field. Exclusive response to pure tones was found in neurons of the rostral dorsal field. Neurons preferring sinusoidal frequency modulations were located in the primary field and the anterior and posterior dorsal fields adjacent to the primary area. Linear frequency modulations optimally activated only neurons of the dorsal part of the dorsal field. Noise-selective neurons were found in the dorsal fields bordering the primary area and the extreme caudal edge of the primary field. The data provide a survey of the functional organization of the horseshoe bat's auditory cortex in real coordinates with the support of cytoarchitectural boundaries and connectional data.

Acoustic Stimulation↗

Neural representation of sound amplitude in the auditory cortex: effects of noise masking.

Single auditory cortical neurons express their sensitivity to the amplitude of a preferred-frequency tone pulse as either a monotonic, saturating intensity profile or as a non-monotonic, bell-shaped intensity function. In the presence of continuous, wideband noise masking, the tone intensity profile is displaced toward higher tone levels. The magnitude of the tone threshold adjustments brought about by increments in noise level very closely match the elevations in noise amplitude. The mechanisms underlying the threshold adjustments likely include neural adaptation. This is because the tone threshold shifts seen in the spike count data are paralleled by spike latency data, and because recovery of tonal sensitivity following noise offset proceeds in a negatively-accelerating fashion. In some instances, the slope of the masked tone intensity profile is greater than that for unmasked tones. For masked tone levels evoking submaximal responses, this has the consequence that cortical responses to masked tones are somewhat more salient than those for unmasked tones of comparable suprathreshold level. These observations bolster our understanding of the psychophysics of noise-masking in normal listeners, and they provide a partial explanation of the difficulty shown by patients with temporal lobe lesions in discriminating signals in noise.

Animals↗

Magnetoencephalographic recordings demonstrate attentional modulation of mismatch-related neural activity in human auditory cortex.

It is widely agreed that the negative brain potential elicited at 150-200 ms by a deviant, less intense sound in a repetitive series can be modulated by attention. To investigate whether this modulation represents a genuine attention effect on the mismatch negativity (MMN) arising from auditory cortex or attention-related activity from another brain region, we recorded both the MMN and the mismatch magnetic field (MMF) elicited by such deviants in a dichotic listening task. Deviant tones in the attended ear elicited a sizable MMF that was well modeled as a dipolar source in auditory cortex. Both the MMN and MMF to unattended-ear deviants were highly attenuated. These findings support the view that the MMN/MMF elicited in auditory cortex by intensity deviants, and thus the underlying feature-analysis and mismatch-detection processes, are not strongly automatic but rather can be gated or suppressed if attention is strongly focused elsewhere.

Acoustic Stimulation↗

Human primary auditory cortex in women and men.

Specific patterns of anatomical symmetry or asymmetry have been associated with sex differences in human brain structure and function. An observer-independent cytoarchitectonic method for the quantification of cell volume densities and areal borders was used to investigate the size and microstructure of primary auditory cortex (Brodmann area 41) in female (n = 14) and male (n = 13) postmortem brains. The total brain volume-adjusted volume of the primary auditory cortex was significantly larger in women than in men bilaterally. Inverse asymmetry towards the right side, as opposed to well-known asymmetries towards the left side, was more frequent in women. Laminar cell volume densities of BA 41 showed no gender effect. The morphometric data confirm (in part) gender differences in the cerebral organization of primary auditory cortex.

Adult↗

Auditory evoked potentials from the primary auditory cortex of the cat: topographic and pharmacological studies.

Wave VI (8.4 msec) of the brain-stem auditory evoked potential (BAEP) was maximal in a discrete region of primary auditory cortex (AI) of the anesthetized cat. Wave VI underwent rapid amplitude decrease over millimeter distances in the AI region and followed high stimulation rates. Wave VI did not show intracortical polarity inversion nor was it abolished by epicortical or intracortical GABA administration. The data are compatible with a wave VI source in the terminal axons of the thalamo-cortical radiations. Middle latency auditory responses (MAEPs) generated 10-40 msec after auditory stimulation were also recorded in a circumscribed area of AI. In contrast to wave VI, these primary auditory cortex potentials (Pa 18.3 msec; Nb 31.9 msec) underwent transcortical polarity inversion, correlated with intracortical multi-unit activity in the AI region and were reversibly altered or abolished by epicortical or intracortical GABA administration to the AI region. The data suggest that the Pa and Nb components of the cat MAEP are intracortically generated by neuronal elements in the AI region.

Acoustic Stimulation↗

Cytoarchitectural and axonal maturation in human auditory cortex.

This study followed the maturation of human auditory cortex from the beginning of the second trimester of gestation to young adulthood. Histological and immunohistochemical techniques were used to trace the development of a laminar cytoarchitecture and an adult pattern of axonal neurofilament expression. From the 16th fetal week to the 4th postnatal month, the cortex progresses from a marginal layer and an undifferentiated cortical plate to incipient lamination. Between the 22nd fetal week and the 4th postnatal month, a two-tiered band of neurofilament-immunoreactive axons develops in layer I, but subsequent to the 4th month, the number of immunopositive axons in this layer is greatly reduced. Between the middle of the first year of life and age 3 years, the laminar pattern of cytoarchitecture becomes fully mature and a network of immunostained axons develops in layers VI, V, IV, and IlIc. This axonal plexus in the deep cortical layers continues to increase in density until age 5. Beginning at 5 years of age, a network of neurofilament-positive axons develops in the superficial layers IIIb, IIIa, and II, and by 11-12 years of age, overall axonal density is equivalent to that seen in young adulthood. This extended time span of axonal maturation has implications for the emergence of auditory cortical function.

Adolescent↗

A chronic microelectrode investigation of the tonotopic organization of human auditory cortex.

We investigated the functional organization of human auditory cortex using a new chronic microelectrode technique. Tonotopic mapping data was obtained at the single unit level for the first time in humans. All sound-driven units were noted to have frequency-dependent response patterns. The majority of units (73%) demonstrated sharply tuned excitatory best-frequency responses. Twenty seven percent of units showed wide receptive fields, representing excitatory responses to almost the entire range of frequencies presented. A tonotopic pattern was observed with best frequencies systematically increasing as more medial-caudal recording sites were sampled.

Acoustic Stimulation↗