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PubMed2004

An investigation of the role of auditory cortex in sound localization using muscimol-releasing Elvax.

Lesion studies suggest that primary auditory cortex (A1) is required for accurate sound localization by carnivores and primates. In order to elucidate further its role in spatial hearing, we examined the behavioural consequences of reversibly inactivating ferret A1 over long periods, using Elvax implants releasing the GABA(A) receptor agonist muscimol. Sub-dural polymer placements were shown to deliver relatively constant levels of muscimol to underlying cortex for >5 months. The measured diffusion of muscimol beneath and around the implant was limited to 1 mm. Cortical silencing was assessed electrophysiologically in both auditory and visual cortices. This exhibited rapid onset and was reversed within a few hours of implant removal. Inactivation of cortical neurons extended to all layers for implants lasting up to 6 weeks and throughout at least layers I-IV for longer placements, whereas thalamic activity in layer IV appeared to be unaffected. Blockade of cortical neurons in the deeper layers was restricted to < or = 500 microm from the edge of the implant, but was usually more widespread in the superficial layers. In contrast, drug-free Elvax implants had little discernible effect on the responses of the underlying cortical neurons. Bilateral implants of muscimol-Elvax over A1 produced significant deficits in the localization of brief sounds in horizontal space and particularly a reduced ability to discriminate between anterior and posterior sound sources. The performance of these ferrets gradually improved over the period in which the Elvax was in place and attained that of control animals following its removal. Although similar in nature, these deficits were less pronounced than those caused by cortical lesions and suggest a specific role for A1 in resolving the spatial ambiguities inherent in auditory localization cues.

Acoustic Stimulation↗
PubMed2005

Auditory plasticity: vocal output shapes auditory cortex.

Studies in humans and songbirds have revealed a close link between vocal output and hearing. Now experiments in marmosets have shown that self-generated vocalizations can modulate the activity of neurons in the auditory cortex and even remodel their response properties.

Animals↗
PubMed2004

Effects of an acute acoustic trauma on the representation of a voice onset time continuum in cat primary auditory cortex.

Here we show that hearing loss associated with an impairment of speech recognition causes a decrease in neural temporal resolution. In order to assess central auditory system changes in temporal resolution, we investigated the effect of an acute hearing loss on the representation of a voice onset time (VOT) and gap-duration continuum in primary auditory cortex (AI) of the ketamine-anesthetized cat. Multiple single-unit activity related to the presentation of a /ba/-/pa/ continuum--in which VOT was varied in 5-ms step from 0 to 70 ms-- was recorded from the same sites before and after an acoustic trauma using two 8-electrode arrays. We also obtained data for gaps, of duration equal to the VOT, embedded in noise 5 ms after the onset. We specifically analyzed the maximum firing rate (FRmax), related to the presentation of the vowel or trailing noise burst, as a function of VOT and gap duration. The changes in FRmax for /ba/-/pa/ continuum as a function of VOT match the psychometric function for categorical perception of /ba/-/pa/ modeled by a sigmoid function. An acoustic trauma made the sigmoid fitting functions shallower, and shifted them toward higher values of VOT. The less steep fitting function may be a neural correlate of an impaired psychoacoustic temporal resolution, because the ambiguity between /ba/ and /pa/ should consequently be increased. The present study is the first one in showing an impairment of the temporal resolution of neurons in AI caused by an acute acoustic trauma.

Acute Disease↗
PubMed2002

Responses of neurons in cat primary auditory cortex to bird chirps: effects of temporal and spectral context.

The responses of neurons to natural sounds and simplified natural sounds were recorded in the primary auditory cortex (AI) of halothane-anesthetized cats. Bird chirps were used as the base natural stimuli. They were first presented within the original acoustic context (at least 250 msec of sounds before and after each chirp). The first simplification step consisted of extracting a short segment containing just the chirp from the longer segment. For the second step, the chirp was cleaned of its accompanying background noise. Finally, each chirp was replaced by an artificial version that had approximately the same frequency trajectory but with constant amplitude. Neurons had a wide range of different response patterns to these stimuli, and many neurons had late response components in addition, or instead of, their onset responses. In general, every simplification step had a substantial influence on the responses. Neither the extracted chirp nor the clean chirp evoked a similar response to the chirp presented within its acoustic context. The extracted chirp evoked different responses than its clean version. The artificial chirps evoked stronger responses with a shorter latency than the corresponding clean chirp because of envelope differences. These results illustrate the sensitivity of neurons in AI to small perturbations of their acoustic input. In particular, they pose a challenge to models based on linear summation of energy within a spectrotemporal receptive field.

Acoustic Stimulation↗
PubMed1999

Functional mapping of transsynaptic effects of local manipulation of inhibition in gerbil auditory cortex.

Cortical networks are under the tonic influence of inhibition which is mainly mediated by GABA. The state of inhibition of small neuronal populations in the auditory cortex (AC) field AI of gerbils was altered by local microinjection of GABA, of the GABA(A)-receptor agonist 4-piperidine-sulfonic acid (P4S) and the GABA(A)-receptor antagonists bicuculline methiodide (BMI) and SR-95531. In order to elucidate direct and transsynaptic effects of the alterations of inhibition produced by these substances we used the 2-fluoro-2-deoxy-D-[(14)C(U)] glucose (FDG) mapping method. The injection of GABA (10 mM) caused no significant changes in FDG labeling but P4S caused a marked decrease of local FDG uptake in a small region surrounding the injection site but in no other region. The injection of the GABA(A)-receptor antagonists caused massive increases of FDG uptake within the entire ipsilateral AC, whereas the contralateral AC was not significantly affected in spite of prominent callosal connections. However, disinhibited excitatory output from the ipsilateral AC is suggested by a strong increase in FDG labeling of the corticothalamic fiber tract and ipsilateral structures like medial geniculate nucleus, caudal striatum, and lateral amygdaloid nucleus and a structure at the caudoventral margin of the thalamic reticular nucleus, presumably the subgeniculate nucleus, a structure with hitherto unknown connections and function. No alteration of FDG uptake could be detected in the inferior colliculus, another main descending target structure of the AC. In summary, the effects resulting from microinjection of GABA(A)-receptor antagonists reflect a differential influence of the AC on its anatomically connected target regions. The findings demonstrate the potential of the method of focal application of neuroactive substances in combination with the FDG technique for mapping their transsynaptic influences which are hard to derive from anatomical tracing studies alone.

Animals↗
PubMed1996

Neuronal assembly dynamics in the rat auditory cortex during reorganization induced by intracortical microstimulation.

Single neurons, acting alone, cannot account for the complex and rapid computations that are routinely accomplished by the behaving nervous system. Recent studies with separable multineuron recordings are showing that neuronal assemblies can indeed be detected and that their organization is very dynamic, depending on variables such as time, physical stimulus, and context. Here we explore both single-neuron and assembly properties in the rat's auditory cortex. Acoustic stimuli are used as a normal, physiological input, and weak electrical intracortical microstimulation (ICMS) as a perturbation that forces a rapid cortical reorganization. In this setting, various aspects of neuronal interactions are changed by the ICMS. We found that cortical neurons exhibited highly synchronous oscillatory firing patterns that were enhanced by ICMS. Cross-correlation studies between two spike trains showed that statistically significant correlations depended on the anatomical distance between the two neurons. ICMS changed the strength and the local number of such correlations. Joint petristimulus analysis and gravity analysis showed that the correlation between neuronal activities varied dynamically at several time scales. We have identified neuronal assemblies in two ways, defined through similarity of receptive field properties and defined through correlated firing. Close anatomical spacing between neurons was conducive to, but not sufficient for membership in, the same assembly with either definition. ICMS changed cortical organization by altering assembly membership. Our data show that neuronal assemblies in the rat auditory cortex can be established transiently in time and that their membership is dynamic.

Acoustic Stimulation↗
PubMed1988

Projections from the auditory cortex to the nucleus of the oculomotor nerve. Silver impregnation and radioautographical study.

A monosynaptic, bilateral connection, between morphophysiological auditory cortical areas and the nucleus of the oculomotor nerve was established by means of contemporary silver impregnation methods and radioautographical technique. Degenerating (resp.--labeled) axons enter the oculomotor nucleus neuropil diffusely from rostro-dorso-laterally, and proceed caudo-ventro-medially. The number of degenerating (resp.--labeled) preterminals and terminals is larger within the territory of the nucleus, located ipsilaterally to the cortical lesion (resp.--isotope injection). The presently described projection is relatively very modest, when compared with the substantial connections of the auditory cortex with the subcortical centers of the auditory system. However, the direct temporooculomotor pathway might serve as a rapid and effective mechanism for the vision-turning reflex after auditory stimulation.

Animals↗
PubMed2001

Sequence dependence of post-tetanic potentiation after sequential heterosynaptic stimulation in the rat auditory cortex.

1. To investigate the mechanisms for the coding stimulus sequence in the auditory cortex (AC), post-tetanic potentiation (PTP) was recorded after sequentially combined heterosynaptic stimulation was applied in rat AC slices. 2. Brief tetanic stimulation (TS) was applied at two sites on AC slices at intervals of 0.5-10 s. PTP of field potentials was induced by the earlier TS, rather than the later TS. PTP was followed by sequence-dependent long-term potentiation (LTP). 3. Using Ca(2+) imaging in the slices loaded with rhod-2, a Ca(2+) indicator, a sequence-dependent distribution of PTP was found in AC slices. 4. The sequence-dependent PTP in excitatory postsynaptic potentials (EPSPs) was observed in supragranular pyramidal neurons. 5. The sequence dependence of PTP was not significantly affected by 1 microM bicuculline, an antagonist of GABA(A) receptors, or 100 microM 2-hydroxysaclofen, an antagonist of GABA(B) receptors. 6. Depolarization and firing recorded in pyramidal neurons during the later TS were less vigorous than when the slices were incubated in the control medium. However, this suppression of the responses during the later TS was not observed in the presence of 50 microM atropine, an antagonist of muscarinic receptors. 7. PTP was induced by the earlier and later TS in the presence of 50 microM atropine, so that the sequence dependence of PTP was abolished. Pirenzepine (50 microM), an antagonist of muscarinic M1 receptors, but not methoctramine (30 microM), an antagonist of M2 receptors, eliminated the sequence dependence of PTP. 8. These findings suggest that the sequence dependence of PTP in AC might have a role in the temporal processing of auditory information on the scale of seconds.

Animals↗
PubMed1988

Delay-tuned combination-sensitive neurons in the auditory cortex of the vocalizing mustached bat.

1. FM-FM neurons in the auditory cortex of the mustached bat are sensitive to a pair of frequency-modulated (FM) sounds that simulates an FM component of the orientation sound and an FM component of the echo. These neurons are tuned to particular delays between the two FM components, suggesting an encoding of target range information. The response properties of these FM-FM neurons, however, have previously been studied only with synthesized orientation sounds and echoes delivered from a loud-speaker as substitutes for the bat's own orientation sounds and corresponding echoes. In this study, the combination sensitivity and delay tuning of FM-FM neurons were examined while the bat was actively vocalizing. 2. When the bat produced orientation sounds in an anechoic environment, or synthesized single FM echoes were delivered to a silent bat, the FM-FM neurons showed weak or no response. In contrast, when synthesized FM echoes were delivered with a particular delay from the FM component of the vocalized orientation sounds, the FM-FM neurons exhibited strong facilitative responses. 3. In both the vocalizing bats and the silent bats with substituted synthesized orientation sounds, all FM-FM neurons tested responded preferentially to the same echo harmonic (FM2, FM3, or FM4). 4. In vocalizing bats, FM-FM neurons showed maximum response to an echo FM component delivered with a particular delay (best delay) from an FM component in the orientation sound. Best delays measured with vocalized orientation sounds were nearly the same as those measured with synthesized orientation sounds. 5. The equivalent effect of a vocalized orientation sound and a synthesized FM1 component on the activity of FM-FM neurons indicates that, during echolocation, the FM1 component in the vocalized orientation sound stimulates the auditory system and conditions the FM-FM neurons to be sensitive to echoes with particular delays from the vocalized orientation sounds. 6. The amount of vocal self-stimulation to the inner ear by the bat's own vocalized sounds was measured by recording cochlear microphonic potentials (CMs). Spectral analysis of CM indicated that the amount of vocal self-stimulation by each harmonic of an orientation sound was equivalent to a sound of 70 dB sound pressure level (SPL) for the first harmonic (H1), 91 dB SPL for H2, 83 dB SPL for H3, and 70 dB SPL for H4, when the amplitude of the vocalized sound was 117 dB SPL at 5 cm in front of the bat's mouth.

Animals↗
PubMed1987

Responses of single neurons in cat auditory cortex to time-varying stimuli: linear amplitude modulations.

Single neurons in the auditory cortex of anesthetized cats were examined quantitatively for their sensitivity to the sound pressure level of characteristic frequency (CF) tone pulses, and to 6 dB, linear modulations in the amplitude of a continuous CF carrier tone. The direction and rate of amplitude modulation (AM), and the carrier level on which it was imposed, were manipulated parametrically. Studied with amplitude modulations, the majority of neurons responded only to intensity increments. The minimum carrier level upon which an amplitude modulation was able to evoke spike discharges was typically comparable to the tone pulse threshold SPL. For many neurons, an "intensity increment response area", i.e., the domain of AM rate and carrier level conjunctions within which a 6 dB AM was able to evoke discharges, could be delimited. For many neurons, preferred rate of AM drifted from high to low with increases in the carrier level on which the modulation was imposed. The most vigorous responses to AM stimuli often occurred when the carrier levels were associated with the rising slope or the peak of the tone pulse rate intensity function. It may be possible to understand the general form of AM response areas in terms of short-term adaptation, the disposition of excitatory and inhibitory tone pulse response areas, and the spectra of the AM stimuli used.

Acoustic Stimulation↗
PubMed1998

Spectral-ripple representation of steady-state vowels in primary auditory cortex.

Responses to various steady-state vowels were recorded in single units in the primary auditory cortex (AI) of the barbiturate-anaesthetized ferret. Six vowels were presented (/a/, /epsilon/, 2 different /i/'s, and 2 different /u/'s) in a natural voiced and a synthetic unvoiced mode. In addition, the responses to broadband stimuli with a sinusoidally shaped spectral envelope (called ripple stimuli) were recorded in each cell, and the response field (RF), which consists of both excitatory and inhibitory regions, was derived from the ripple transfer function. We examined whether the vowel responses could be predicted using a linear ripple analysis method [Shamma et al., Auditory Neurosci. 1, 233-254 (1995)], i.e., by cross correlating the RF of the single unit, and the smoothed spectral envelope of the vowel. We found that for most AI cells (71%) the relative responses to natural vowels could be predicted on the basis of this method. Responses and prediction results for unvoiced and voiced vowels were very similar, suggesting that the spectral fine structure may not play a significant role in the neuron's response to the vowels. Predictions on the basis of the entire RF were significantly better than based solely on best frequency (BF) (or "place"). These findings confirm the ripple analysis method as a valid method to characterize AI responses to broadband sounds as we proposed in a previous paper using synthesized spectra [Shamma and Versnel, Auditory Neurosci. 1, 255-270 (1995)].

Animals↗
PubMed1985

Sensitivity of cat primary auditory cortex (AI) neurons to the direction and rate of frequency modulation.

Responses of 65 single auditory cortex (AI) neurons to frequency-modulated (FM) sweeps with different rates and direction of frequency change were examined quantitatively. Most units responded differentially depending on the characteristics of the FM sweep stimulus. Sixty-five percent of the units encountered responded at least twice as well for one direction of the FM sweep as for the other direction. Of these direction selective neurons, 67% preferred downward-directed FM sweeps (i.e. changing from high to low frequencies) while only 33% preferred upward-directed FM sweeps. The preference for downward-directed FM sweeps was especially clear in EI cells. In addition, cortical neurons often displayed sensitivity to the rate of frequency modulation (speed sensitivity).

Acoustic Stimulation↗
PubMed1999

Origins of medial geniculate body projections to physiologically defined zones of rat primary auditory cortex.

Medial geniculate body neurons projecting to physiologically identified subregions of rat primary auditory cortex (area 41, Te1) were labeled with horseradish peroxidase in adult rats. The goals were to determine the type(s) of projection neuron and the spatial arrangement of these cells with respect to thalamic subdivisions. Maps of best frequency were made with single neuron or unit cluster extracellular recording at depths of 500-800 microm, which correspond to layers III-IV in Nissl preparations. Tracer injections were made in different cortical isofrequency regions (2, 11, 22, or 38 kHz, respectively). Labeled neurons were plotted on representative sections upon which the architectonic subdivisions were drawn independently. Most of the cells of origin lay in the ventral division in every experiment. Injections at low frequencies labeled bands of neurons laterally in the ventral division; progressively more rostral deposits at higher frequencies labeled bands or clusters more medially in the ventral division, and through most of its caudo-rostral extent. Medial division labeling was variable. Labeled cells were always in the lateral half of the nucleus and were often scattered. There were few labeled cells in the dorsal division. Seven types of thalamocortical neuron were identified: ventral division cells had a tufted branching pattern, while medial division neurons have heterogeneous shapes and sizes and were larger. Dorsal division neurons had a radiate branching pattern. The size range of labeled neurons spanned that of Nissl stained neuronal somata. Area 41 may receive two types of thalamic projection: ventral division input is strongly convergent, highly topographic, spatially focal, and restricted to one type of neuron only, while the medial division projection is more divergent, coarsely topographical, involves multiple cortical areas, and has several varieties of projection neuron. Despite species differences in local circuitry, many facets of thalamocortical organization are conserved in phylogeny.

Animals↗
PubMed1993

Spatiotemporal observation of guinea pig auditory cortex with optical recording.

The spatiotemporal characteristics of neural activity in the guinea pig auditory cortex were studied in order to clarify neural processing and coding mechanisms of complex sounds. We used a multi-channel optical recording system for a voltage-sensitive dye: RH795. The experimental results showed that a boomerang-shaped moving pattern of optical response appeared on the cortical surface in response to complex sounds (clicks) stimulation, but a rather fixed pattern appeared in response to tone burst stimulation. The tonotopical organization observed using microelectrode was not directly visible, but a similar topographic pattern was evidenced by selecting regions of a strong response evoked by tone-bursts. The correlative functions of regionwide responses may indicate a parallel and serial neural processing structure.

Acoustic Stimulation↗