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PubMed1986

Activation of the auditory cortex by cochlear stimulation in a deaf patient.

We have recorded cerebral electric and magnetic responses to electric stimulation of the auditory nerve in a deaf patient with multichannel cochlear prosthesis. The electric response peaked with a vertex-negative deflection at 65 ms. Clear magnetic responses, coinciding with the vertex potential, were obtained only over the hemisphere ipsilateral to the stimulation; the field pattern indicated activation of the auditory cortex within the Sylvian fissure. The results suggest modification of the central auditory pathways in this patient deaf from early childhood.

Auditory Cortex↗
PubMed1976

Relation of unit spike discharges and evoked potentials in the auditory cortex of cats.

Extracellular microelectrode recordings were made from the auditory cortex of anaesthetized cats during acoustic click stimulation. The microelectrode of low resistance allowed to record evoked field potentials and unit discharges simultaneously. In distant extracellular leads the relation of unit discharges and field potentials was equivocal. Near extracellular leads revealed that the antidromic invasion of the somadendritic membrane by excitation is a frequency dependent process (just as evoked field potentials) while spike potentials can reliably be elicited from the initial segment at high frequencies. It is assumed that the excitation spreading from the initial segment to the soma-dendritic membrane represents an important component of the evoked potentials, and their frequency dependence may be traced back to inhibitions activated by afferent impulses.

Acoustic Stimulation↗
PubMed1993

Neurons in the inferior colliculus, auditory cortex and pontine nuclei of the FM bat, Eptesicus fucus respond to pulse repetition rate differently.

Single-neuron responses to pulse repetition rate in the inferior colliculus, auditory cortex and pontine nuclei of the FM bat, Eptesicus fuscus were studied under free-field stimulation conditions. The best frequency (BF) and minimum threshold (MT) of each neuron were first determined with a 4 ms pulse broadcast from a specific point (response center) of the bat's frontal auditory space at which the neuron had maximal spatial sensitivity. The neuron's intensity-rate function was then studied with a 4 ms BF pulse delivered at 10 dB increments above its MT in order to determine the best intensity to which the neuron discharged maximally. The neuron's discharge pattern and number of impulses to 32 trials of 300 ms train stimuli, which consisted of different number of 4 ms BF and best intensity pulses (1, 2, 3, 8, 12, 19, 24, 29 pulses/train) and delivered at an interpulse interval of 1000, 250, 170, 100, 40, 25, 15, 12 and 10 ms (i.e. at a pulse repetition rate of 1, 4, 6, 10, 25, 40, 67, 83, 100 pulses/s), were sequentially recorded. All neurons recorded from the inferior colliculus, auditory cortex and pontine nuclei discharged phasically (1-3 impulses) but they responded to the pulse repetition rate in different manners. More than 63% of 38 inferior collicular and 65 pontine neurons studied discharged impulses to each pulse within a train stimulus when the pulse repetition rate was up to 40 pulses/s.(ABSTRACT TRUNCATED AT 250 WORDS)

Acoustic Stimulation↗
PubMed2004

Auditory stream segregation in monkey auditory cortex: effects of frequency separation, presentation rate, and tone duration.

Auditory stream segregation refers to the organization of sequential sounds into "perceptual streams" reflecting individual environmental sound sources. In the present study, sequences of alternating high and low tones, "...ABAB...," similar to those used in psychoacoustic experiments on stream segregation, were presented to awake monkeys while neural activity was recorded in primary auditory cortex (A1). Tone frequency separation (AF), tone presentation rate (PR), and tone duration (TD) were systematically varied to examine whether neural responses correlate with effects of these variables on perceptual stream segregation. "A" tones were fixed at the best frequency of the recording site, while "B" tones were displaced in frequency from "A" tones by an amount = delta F. As PR increased, "B" tone responses decreased in amplitude to a greater extent than "A" tone responses, yielding neural response patterns dominated by "A" tone responses occurring at half the alternation rate. Increasing TD facilitated the differential attenuation of "B" tone responses. These findings parallel psychoacoustic data and suggest a physiological model of stream segregation whereby increasing delta F, PR, or TD enhances spatial differentiation of "A" tone and "B" tone responses along the tonotopic map in A1.

Acoustic Stimulation↗
PubMed2004

Intracortical pathways determine breadth of subthreshold frequency receptive fields in primary auditory cortex.

To examine the basis of frequency receptive fields in auditory cortex (ACx), we have recorded intracellular (whole cell) and extracellular (local field potential, LFP) responses to tones in anesthetized rats. Frequency receptive fields derived from excitatory postsynaptic potentials (EPSPs) and LFPs from the same location resembled each other in terms of characteristic frequency (CF) and breadth of tuning, suggesting that LFPs reflect local synaptic (including subthreshold) activity. Subthreshold EPSP and LFP receptive fields were remarkably broad, often spanning five octaves (the maximum tested) at moderate intensities (40-50 dB above threshold). To identify receptive-field features that are generated intracortically, we microinjected the GABA(A) receptor agonist muscimol (0.2-5.1 mM, 1-5 microl) into ACx. Muscimol dramatically reduced LFP amplitude and reduced receptive-field bandwidth, implicating intracortical contributions to these features but had lesser effects on CF response threshold or onset latency, suggesting minimal loss of thalamocortical input. Reversal of muscimol's inhibition preferentially at the recording site by diffusion from the recording pipette of the GABA(A) receptor antagonist picrotoxin (0.01-100 microM) disinhibited responses to CF stimuli more than responses to spectrally distant, non-CF stimuli. We propose that thalamocortical and intracortical pathways preferentially contribute to responses evoked by CF and non-CF stimuli, respectively, and that intracortical projections linking frequency representations determine the breadth of receptive fields in primary ACx. Broad, subthreshold receptive fields may distinguish ACx from subcortical auditory relay nuclei, promote integrated responses to spectrotemporally complex stimuli, and provide a substrate for plasticity of cortical receptive fields and maps.

Acoustic Stimulation↗
PubMed1999

Basal forebrain stimulation induces discriminative receptive field plasticity in the auditory cortex.

Learning alters receptive field (RF) tuning in the primary auditory cortex (ACx) to emphasize the frequency of a tonal conditioned stimulus. RF plasticity is a candidate substrate of memory, as it is associative, specific, discriminative, rapidly induced, and enduring. The authors hypothesized that it is produced by the release of acetylcholine in the ACx from the basal forebrain (BasF), caused by presentation of reinforced but not nonreinforced conditioned stimuli. Waking adult male Hartley guinea pigs (n = 16) received 1 of 2 tones followed by BasF stimulation, in a single session of 30 pseudo-random order trials each. RFs from neuronal discharges before and after differential pairing revealed the induction of predicted plasticity, as well as increased responses to the paired tone and decreased responses to the unpaired tone. Thus, highly specific, learning-induced RF plasticity in the ACx may be produced by activation of the BasF by a reinforced conditioned stimulus.

Acoustic Stimulation↗
PubMed2000

Voice-selective areas in human auditory cortex.

The human voice contains in its acoustic structure a wealth of information on the speaker's identity and emotional state which we perceive with remarkable ease and accuracy. Although the perception of speaker-related features of voice plays a major role in human communication, little is known about its neural basis. Here we show, using functional magnetic resonance imaging in human volunteers, that voice-selective regions can be found bilaterally along the upper bank of the superior temporal sulcus (STS). These regions showed greater neuronal activity when subjects listened passively to vocal sounds, whether speech or non-speech, than to non-vocal environmental sounds. Central STS regions also displayed a high degree of selectivity by responding significantly more to vocal sounds than to matched control stimuli, including scrambled voices and amplitude-modulated noise. Moreover, their response to stimuli degraded by frequency filtering paralleled the subjects' behavioural performance in voice-perception tasks that used these stimuli. The voice-selective areas in the STS may represent the counterpart of the face-selective areas in human visual cortex; their existence sheds new light on the functional architecture of the human auditory cortex.

Acoustic Stimulation↗
PubMed1997

Positional, directional and speed selectivities in the primary auditory cortex of the cat.

Responses of high-frequency primary auditory cortex (A1) neurons of the cat to noise stimulation were obtained in a quasianechoic chamber using a static and an apparently moving stimulus presented at similar azimuths. Simulated motion toward right or left as well as different simulated velocities were used. Under static stimulation, most units were contralateral-preferring followed by ipsilateral- and midline-preferring. Some were omnidirectional and a few were unclassifiable. Width of tuning was similar for contralateral-, ipsilateral- and midline-preferring units. Overall, about 25% were finely tuned (< 20 degrees) and the remaining were broadly tuned (> or =20 degrees). All cells sampled with static stimulation responded to apparent motion. About one quarter of the units were sensitive to the direction of the simulated moving noise in that they responded at least twice as much to one direction as to the other. Almost all directional contralateral-preferring units responded more when the apparent motion was directed toward ipsilateral azimuths, whereas all directional ipsilateral-preferring units responded preferentially to contralaterally oriented motion. In some units, up to five apparent speeds were tested. About half the units were not speed-selective (46%). The other cells were tuned to a preferential speed (40%), decreased their response as the apparent speed increased (10%) or displayed direction-dependent speed selectivity (4%). These results indicate that moving-sound sources are processed by some A1 single units.

Acoustic Stimulation↗
PubMed2005

An oscillatory hierarchy controlling neuronal excitability and stimulus processing in the auditory cortex.

EEG oscillations are hypothesized to reflect cyclical variations in the neuronal excitability, with particular frequency bands reflecting differing spatial scales of brain operation. However, despite decades of clinical and scientific investigation, there is no unifying theory of EEG organization, and the role of ongoing activity in sensory processing remains controversial. This study analyzed laminar profiles of synaptic activity [current source density CSD] and multiunit activity (MUA), both spontaneous and stimulus-driven, in primary auditory cortex of awake macaque monkeys. Our results reveal that the EEG is hierarchically organized; delta (1-4 Hz) phase modulates theta (4-10 Hz) amplitude, and theta phase modulates gamma (30-50 Hz) amplitude. This oscillatory hierarchy controls baseline excitability and thus stimulus-related responses in a neuronal ensemble. We propose that the hierarchical organization of ambient oscillatory activity allows auditory cortex to structure its temporal activity pattern so as to optimize the processing of rhythmic inputs.

Acoustic Stimulation↗
PubMedDate not supplied

An animal model of auditory cortex prostheses.

Experiments aimed at rehabilitating deaf and blind patients with cortical prostheses were first conducted decades ago, but epicortical electrodes allowed only crude information transfer. Here we report that in Mongolian gerbils with electrodes implanted in input layers of the primary auditory cortex, spatial, temporal and spatiotemporal variations in intracortical stimulation all lead to perceptual differences as evidenced by discrimination training. For some stimulus regimes discrimination learning was as fast as with intracochlear stimulation in this animal. Intracortical stimulation induced field potentials and 2-deoxyglucose labeling patterns in primary auditory cortex similar to those induced by auditory click or tone stimuli, respectively. Given the common organization principles of neocortical areas, these results are presumably also of significance to prostheses interfacing with visual cortex.

Animals↗
PubMed2006

Thalamic connections of the auditory cortex in marmoset monkeys: core and medial belt regions.

In this study and its companion, the cortical and subcortical connections of the medial belt region of the marmoset monkey auditory cortex were compared with the core region. The main objective was to document anatomical features that account for functional differences observed between areas. Injections of retrograde and bi-directional anatomical tracers targeted two core areas (A1 and R), and two medial belt areas (rostromedial [RM] and caudomedial [CM]). Topographically distinct patterns of connections were revealed among subdivisions of the medial geniculate complex (MGC) and multisensory thalamic nuclei, including the suprageniculate (Sg), limitans (Lim), medial pulvinar (PM), and posterior nucleus (Po). The dominant thalamic projection to the CM was the anterior dorsal division (MGad) of the MGC, whereas the posterior dorsal division (MGpd) targeted RM. CM also had substantial input from multisensory nuclei, especially the magnocellular division (MGm) of the MGC. RM had weak multisensory connections. Corticotectal projections of both RM and CM targeted the dorsomedial quadrant of the inferior colliculus, whereas the CM projection also included a pericentral extension around the ventromedial and lateral portion of the central nucleus. Areas A1 and R were characterized by focal topographic connections within the ventral division (MGv) of the MGC, reflecting the tonotopic organization of both core areas. The results indicate that parallel subcortical pathways target the core and medial belt regions and that RM and CM represent functionally distinct areas within the medial belt auditory cortex.

Animals↗
PubMed2001

Role of mammalian auditory cortex in the perception of elementary sound properties.

Studies in several mammalian species have demonstrated that bilateral ablations of the auditory cortex have little effect on simple sound intensity and frequency-based behaviors. In the rat, for example, early experiments have shown that auditory ablations result in virtually no effect on the rat's ability to either detect tones or discriminate frequencies. Such lesion experiments, however, typically examine an animal's performance some time after recovery from ablation surgery. As such, they demonstrate that the cortex is not essential for simple auditory behaviors in the long run. Our study further explores the role of cortex in basic auditory perception by examining whether the cortex is normally involved in these behaviors. In these experiments we reversibly inactivated the rat primary auditory cortex (AI) using the GABA agonist muscimol, while the animals performed a simple auditory task. At the same time we monitored the rat's auditory activity by recording auditory evoked potentials (AEP) from the cortical surface. In contrast to lesion studies, the rapid time course of these experimental conditions preclude reorganization of the auditory system that might otherwise compensate for the loss of cortical processing. Soon after bilateral muscimol application to their AI region, our rats exhibited an acute and profound inability to detect tones. After a few hours this state was followed by a gradual recovery of normal hearing, first of tone detection and, much later, of the ability to discriminate frequencies. Surface muscimol application, at the same time, drastically altered the normal rat AEP. Some of the normal AEP components vanished nearly instantaneously to unveil an underlying waveform, whose size was related to the severity of accompanying behavioral deficits. These results strongly suggest that the cortex is directly involved in basic acoustic processing. Along with observations from accompanying multiunit experiments that related the AEP to AI neuronal activity, our results suggest that a critical amount of activity in the auditory cortex is necessary for normal hearing. It is likely that the involvement of the cortex in simple auditory perceptions has hitherto not been clearly understood because of underlying recovery processes that, in the long-term, safeguard fundamental auditory abilities after cortical injury.

Animals↗
PubMed2005

Primary auditory cortex activation by visual speech: an fMRI study at 3 T.

Recent studies have yielded contradictory evidence on whether visual speech perception (watching articulatory gestures) can activate the human primary auditory cortex. To circumvent confounds due to inter-individual anatomical variation, we defined our subjects' Heschl's gyri and assessed blood oxygenation-dependent signal changes at 3 T within this confined region during visual speech perception and observation of moving circles. Visual speech perception activated Heschl's gyri in nine subjects, with activation in seven of them extending to the area of primary auditory cortex. Activation was significantly stronger during visual speech perception than during observation of the moving circles. Further, a significant hemisphere by stimulus interaction occurred, suggesting left Heschl's gyrus specialization for visual speech processing.

Adult↗
PubMed2001

Linear processing of spatial cues in primary auditory cortex.

To determine the direction of a sound source in space, animals must process a variety of auditory spatial cues, including interaural level and time differences, as well as changes in the sound spectrum caused by the direction-dependent filtering of sound by the outer ear. Behavioural deficits observed when primary auditory cortex (A1) is damaged have led to the widespread view that A1 may have an essential role in this complex computational task. Here we show, however, that the spatial selectivity exhibited by the large majority of A1 neurons is well predicted by a simple linear model, which assumes that neurons additively integrate sound levels in each frequency band and ear. The success of this linear model is surprising, given that computing sound source direction is a necessarily nonlinear operation. However, because linear operations preserve information, our results are consistent with the hypothesis that A1 may also form a gateway to higher, more specialized cortical areas.

Animals↗
PubMed1993

Temporal integration and oscillatory responses of the human auditory cortex revealed by evoked magnetic fields to click trains.

We recorded neuromagnetic evoked responses from the right auditory cortex of 7 healthy adults with a 24-channel planar SQUID gradiometer. The stimuli were 200-ms click trains presented at rates of 40, 80, 160 and 320 Hz, with interstimulus intervals (ISIs) of 1 and 4 s. The transient N100m response to the train onset depended on the click rate: the peak latency shortened to the same extent as the interval between successive clicks decreased in trains with rates from 40 Hz to 320 Hz. The N100m amplitude increased simultaneously, saturating at rates of 160-320 Hz. The mean N100m latency was slightly longer with the 1-s than with the 4-s ISI for all click rates. The systematic changes of the N100m amplitude and latency according to click rate demonstrate the importance of temporal integration for N100m generation, and imply an integration time of 20-25 ms. The 20- and 40-Hz click trains also elicited oscillatory 40-Hz responses 80-250 ms after the train onset. The 40-Hz responses were more resistant than N100m to changes of the ISI, and their sources slightly differed from those of N100m. These two responses evidently reflect different aspects of auditory processing.

Acoustic Stimulation↗
PubMed1996

Modulation of cerebral blood flow in the human auditory cortex during speech: role of motor-to-sensory discharges.

To investigate mechanisms of audio-vocal interactions in the human brain, we studied the effect of speech output on modulation of neuronal activity in the auditory cortex. The modulation was assessed indirectly by measuring changes in cerebral blood flow (CBF) during unvoiced speech (whispering). Using positron emission tomography (PET), CBF was measured in eight volunteers as they uttered syllables at each of seven rates (30, 50, 70, 90, 110, 130 or 150/min) during each of the seven 60-s PET scans. Low-intensity white noise was used throughout scanning to mask auditory input contingent on the whispering. We found that, as a function of the increasing syllable rate, CBF increased in the left primary face area, the upper pons, the left planum temporale and the left posterior perisylvian cortex. The latter two regions contain secondary auditory cortex and previously have been implicated in the processing of speech sounds. We conclude that, in the absence of speech-contingent auditory input, the modulation of CBF in the auditory cortex is mediated by motor-to-sensory discharges. As such, it extends our previous findings of oculomotor corollary discharges to the audio-vocal domain.

Adult↗
PubMed2005

Enhanced contrast sensitivity in auditory cortex as cats learn to discriminate sound frequencies.

To better understand the nature and time course for learning-induced cortical reorganization, we examined frequency-specific changes in auditory cortex as cats gradually improved at a difficult sound frequency discrimination task. Three adult cats were trained to discriminate between a tone pip at a fixed target frequency (S-) and a higher deviant frequency (S+). An adaptive training schedule led to an efficient estimate of the frequency discrimination threshold (FDT), which was used to track daily performance. Each cat was also implanted with an array of microwires in auditory cortex. Tone pips with different frequency and amplitude were used to map receptive fields. Onset responses were correlated with training time and the cat's ability to discriminate frequencies. Although lifetime of the neural implants varied among cats, each provided sufficient neural recording to relate at least 3 weeks of learning to response changes in the cortex. An improved FDT was associated with a differential decrease in response strength between the S- frequency and S+ frequencies. Response to the training frequencies gradually located in a local minimum compared to adjacent frequencies (p < 0.001, Cohen's d=0.50). Cortical changes were consistent with a theory of bimodal generalization that enhances stimulus classification by reducing similarity between reinforced and nonreinforced stimuli. Such a strategy may be especially appropriate during an early stage of learning to discriminate similar sounds and differ from later strategies required for fine discrimination.

Animals↗
PubMed1997

The columnar and layer-specific response properties of neurons in the primary auditory cortex of Mongolian gerbils.

The columnar and layer-specific response properties of neurons in the primary auditory cortex (AI) of Mongolian gerbils were studied using single-unit recordings of responses to tone-burst stimuli presented to the ear contralateral to the recording side. During near-radial microelectrode penetrations of the AI in 100-microm steps, the best frequency (BF), best threshold (BT), best amplitude (BA), latency, tuning curve and Q10dB were recorded. Neurons encountered during single penetrations showed similar BFs, indicating a columnar frequency organization, but their latencies and Q10dBs differed. The BAs and BTs recorded within single penetrations often showed a similar value in the middle cortical layers. The latencies and Q10dBs of these neurons exhibited a tendency toward a layer-specific distribution. The latencies of neurons located in layers I-V were longer than those located in layer VI. The Q10dBs of neurons located in layers III and IV were higher than those located in layers I and VI. These results are almost consistent with those of previous studies on frequency representation, and indicated the existence of an integrative mechanism of frequency processing in the AI. This is the first study in which a layer-specific, partially columnar organization for stimulus amplitude is described.

Acoustic Stimulation↗