Evidence for a tonotopic organization of the auditory cortex observed with auditory evoked potentials.
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Acoustic Stimulation↗
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The Organization of Projections from Subdivisions of the Auditory Cortex and Thalamus to the Auditory Sector of the Thalamic Reticular Nucleus in Galago.
Anterograde and retrograde transport techniques were used to study the connexions between different subdivisions of the auditory cortex and thalamus with the thalamic reticular nucleus in the prosimian, Galago. In particular, the goal was to determine whether the primary auditory nucleus, GMv, and its cortical target, area I of the auditory cortex (A I), project to a different region of the auditory sector of the reticular nucleus from the secondary auditory nuclei, GMmc and Po and their cortical targets outside A I. The results show that the projections to and from the auditory sector are indeed segregated: injections of wheatgerm agglutinin-conjugated horseradish peroxidase into either GMmc or Po labelled cells and terminals along the medial, lateral and ventral borders of the auditory sector, forming a U-shaped pattern. Projections from area II of the auditory cortex produced almost an identical pattern of the terminal labelling in the auditory sector. In contrast, injections into GMv-labelled cells and terminals in the centre region of the auditory sector, in the 'interior' of the U-shaped region. Projections from A I were distributed to both the U-shaped border region and the central core of the auditory sector probably because A I received projections from GMmc, Po and GMv. The significance of these results depends on a comparison between the auditory and visual sectors of the reticular nucleus. Both sectors are divided into tiers or subsectors-one related to the primary relay nucleus, i.e. GLd or GMv, and the other related to the secondary relay nuclei, i.e. pulvinar nucleus, GMmc, Po, etc.
Journal Article↗
Response of the primary auditory cortex to electrical stimulation of the auditory nerve in the congenitally deaf white cat.
Neural activity plays an important role in the development and maintenance of sensory pathways. However, while there is considerable experience using cochlear implants in both congenitally deaf adults and children, little is known of the effects of a hearing loss on the development of the auditory cortex. In the present study, cortical evoked potentials, field potentials, and multi- and single-unit activity evoked by electrical stimulation of the auditory nerve were used to study the functional organisation of the auditory cortex in the adult congenitally deaf white cat. The absence of click-evoked auditory brainstem responses during the first weeks of life demonstrated that these animals had no auditory experience. Under barbiturate anaesthesia, cortical potentials could be recorded from the contralateral auditory cortex in response to bipolar electrical stimulation of the cochlea in spite of total auditory deprivation. Threshold, morphology and latency of the evoked potentials varied with the location of the recording electrode, with response latency varying from 10 to 20 ms. There was evidence of threshold shifts with site of the cochlear stimulation in accordance with the known cochleotopic organisation of AI. Thresholds also varied with the configuration of the stimulating electrodes in accordance with changes previously observed in normal hearing animals. Single-unit recordings exhibited properties similar to the evoked potentials. Increasing stimulus intensity resulted in an increase in spike rate and a decrease in latency to a minimum of approximately 8 ms, consistent with latencies recorded in AI of previously normal animals (Raggio and Schreiner, 1994). Single-unit thresholds also varied with the configuration of the stimulating electrodes. Strongly driven responses were followed by a suppression of spontaneous activity. Even at saturation intensities the degree of synchronisation was less than observed when recording from auditory brainstem nuclei. Taken together, in these auditory deprived animals basic response properties of the auditory cortex of the congenitally deaf white cat appear similar to those reported in normal hearing animals in response to electrical stimulation of the auditory nerve. In addition, it seems that the auditory cortex retains at least some rudimentary level of cochleotopic organisation.
Adult↗
Neurons in cat primary auditory cortex sensitive to correlates of auditory motion in three-dimensional space.
Amplitude modulation at the receiver's ears is a characteristic of moving sound sources. When a sound source moves from side to side, stimulus intensity decreases in one ear and increases in the other. When a sound source moves toward or away from the organism, the two ears receive correlated increases or decreases in sound level. We recorded from single cells in the auditory cortex while presenting amplitude modulated pure tones to the two ears which stimulated motion either toward or away from the organism, or from side to side. Our results indicate that auditory cortex neurons can be highly sensitive to these correlates of auditory motion in three dimensional space. Three major classes of neurons were encountered. These included 1) neurons sensitive to azimuthal stimulus motion, 2) neurons sensitive to motion directly toward or away from the organism, and 3) monaural-like neurons. More toward-preferring neurons than away-preferring neurons were encountered, and more units preferred contralateral-directed than ipsilateral-directed movement. The different classes of direction-selective neurons were spatially segregated from each other within the cortex and appear to occur in columns. In addition to their selectivity for different directions of simulated sound source motion, auditory cortex neurons could also be highly selective to AM ramp rate and excursion; these are correlates of sound source velocity.
Acoustic Stimulation↗
Tonotopic organization of the human auditory cortex revealed by transient auditory evoked magnetic fields.
The tonotopic organization of the human auditory cortex has been investigated by systematic measurements of magnetic fields evoked by tone-bursts with carrier frequencies of 250, 500, 1000, 2000 and 4000 Hz. The measured field distribution changes with both time elapsed since stimulus onset and frequency of the stimulus. Nevertheless, the field distribution has always the same overall features and can be approximated by that of an equivalent current dipole located in a semi-infinite volume. This model can be described in terms of 5 parameter values: 3 orthogonal coordinates specifying the dipole location, and amplitude and angle of the dipole moment. The amplitude of the dipole moment is maximal at about 100 msec ('component 100m') and 160 msec ('component 160m') after stimulus onset. The depth estimated for the generator site of the 100m component shows a logarithmic dependence on test frequency whereas no similar behaviour could be observed for the 160m component. Anatomical studies performed in cadaver heads suggest that the equivalent current dipoles of both the 100m and the 160m component are located in the transverse temporal gyri.
Adult↗
Function of dog's auditory cortex in tests involving auditory location cues and directional instrumental response.
Twenty one dogs, distributed into four groups, were trained pre-operatively in differentiation of auditory location or frequency cues. In each group instrumental responses, reinforced by food consisted in placing by the animal its right paw on the side levers. The first differentiation, go-left, go-right task with two location cues, required the animal to place its paw on the lever opposite to the source of the cue. The second differentiation task with the same location cues required placing the paw on the lever located close to the cue. The third task, involving 700 Hz vs. 1,000 Hz tone, required responding to one lever to the presentation of one tone and responding to the opposite lever to the presentation of the other. The last task was a symmetrically reinforced go, no-go differentiation with, again, auditory location cues: the animals were trained to place the paw on a lever to one location cue and to withhold this response to the other location cue. Bilateral ablation of the primary auditory cortex produced a considerable impairment of the performance of the two go-left, go-right tasks involving location cues. The go-left, go-right task employing frequency cues, and the symmetrically reinforced go, no-go task with location cues, were only slightly disturbed by this lesion.
Animals↗
Binaural interactions of single neurons in posterior field of cat auditory cortex.
In the auditory cortex of barbiturate-anesthetized cats, the posterior auditory field (area P) was identified by its tonotopic organization, and single neurons in that field were studied quantitatively with regard to their binaural interactions at their respective best frequencies, using calibrated, sealed stimulating systems. Almost 60% of the neurons studied displayed " summative " binaural interactions in that their responses to binaural, equally intense stimulation of the two ears were stronger than were their responses to monaural stimuli of the same intensity. For these neurons, latent periods were shorter for binaural stimuli than for monaural stimuli. Some field P neurons were sensitive to interaural intensity disparities and manifested that sensitivity in one of two forms. Cells that were excited by stimulation of one ear and inhibited by stimulation of the other typically displayed a sigmoidal relation of spike count to intensive disparity, with spike counts being larger when the disparity favored the contralateral ear. Cells that were unresponsive to monaural stimuli but responded securely to binaural stimuli usually displayed a peaked, nonmonotonic relation of spike count to interaural intensity disparity, with maximal responses being elicited by stimuli with zero or near-zero disparity. Some neurons of low best frequency were sensitive to variations in interaural phase delay. In all cases, this sensitivity was manifested as a cyclical relation of spike count to interaural delay, with the period of the cycle being that of the stimulating tone. The fact that the binaural interactions of field P neurons were similar to those of cells in the primary auditory cortex suggests that the previously described heightened spectral-amplitude selectivity of field P neurons has been achieved without cost to their sensitivity to a variety of parameters of binaural stimulation. The particular sensitivity of cortical neurons to variations in interaural disparities associated with midline or near-midline azimuths might constitute a neural mechanism for the behavioral finding that animals and humans show their greatest acuity in sound localization for stimulus locations in or near the midsagittal plane.
Animals↗
Early chronic blockade of NR2B subunits and transient activation of NMDA receptors modulate LTP in mouse auditory cortex.
In the auditory cortex, the properties of NMDA receptors depend primarily on the ratio of NR2A and NR2B subunits. NR2B subunit expression is high at the beginning of critical period and lower in adulthood. Because NMDA receptors are crucial in triggering long-term potentiation (LTP) and long-term depression, developmental or experience-dependent modification of NMDAR subunit composition is likely to influence synaptic plasticity. To examine how NMDA subunit change during postnatal development affect the adult synaptic plasticity, we employed chronic ifenprodil blockade of NR2B subunits and analyzed evoked field potentials in adult C57BL/6 mice auditory cortex (AC). We found that chronic loss of NR2B activity led to a decline in LTP magnitude in the AC of adult mice. Adding NMDA to the artificial cerebrospinal fluid (ACSF) in blocked mice had the opposite effect, producing LTP magnitudes at or exceeding those found in treated or untreated animals. These results suggest that, even in adulthood when NR2B expression is downregulated, these receptor subunits play an important role in experience-dependent plasticity of mouse auditory cortex. Blockade from P60 did not result in any decrease of LTP amplitude, suggesting that chronic block in postnatal period may permanently affect cortical circuits so that they cannot produce significant LTP in adulthood.
Analysis of Variance↗
Location coding by opponent neural populations in the auditory cortex.
Although the auditory cortex plays a necessary role in sound localization, physiological investigations in the cortex reveal inhomogeneous sampling of auditory space that is difficult to reconcile with localization behavior under the assumption of local spatial coding. Most neurons respond maximally to sounds located far to the left or right side, with few neurons tuned to the frontal midline. Paradoxically, psychophysical studies show optimal spatial acuity across the frontal midline. In this paper, we revisit the problem of inhomogeneous spatial sampling in three fields of cat auditory cortex. In each field, we confirm that neural responses tend to be greatest for lateral positions, but show the greatest modulation for near-midline source locations. Moreover, identification of source locations based on cortical responses shows sharp discrimination of left from right but relatively inaccurate discrimination of locations within each half of space. Motivated by these findings, we explore an opponent-process theory in which sound-source locations are represented by differences in the activity of two broadly tuned channels formed by contra- and ipsilaterally preferring neurons. Finally, we demonstrate a simple model, based on spike-count differences across cortical populations, that provides bias-free, level-invariant localization-and thus also a solution to the "binding problem" of associating spatial information with other nonspatial attributes of sounds.
Acoustic Stimulation↗
Background sounds contribute to spectrotemporal plasticity in primary auditory cortex.
The mammalian auditory system evolved to extract meaningful information from complex acoustic environments. Spectrotemporal selectivity of auditory neurons provides a potential mechanism to represent natural sounds. Experience-dependent plasticity mechanisms can remodel the spectrotemporal selectivity of neurons in primary auditory cortex (A1). Electrical stimulation of the cholinergic nucleus basalis (NB) enables plasticity in A1 that parallels natural learning and is specific to acoustic features associated with NB activity. In this study, we used NB stimulation to explore how cortical networks reorganize after experience with frequency-modulated (FM) sweeps, and how background stimuli contribute to spectrotemporal plasticity in rat auditory cortex. Pairing an 8-4 kHz FM sweep with NB stimulation 300 times per day for 20 days decreased tone thresholds, frequency selectivity, and response latency of A1 neurons in the region of the tonotopic map activated by the sound. In an attempt to modify neuronal response properties across all of A1 the same NB activation was paired in a second group of rats with five downward FM sweeps, each spanning a different octave. No changes in FM selectivity or receptive field (RF) structure were observed when the neural activation was distributed across the cortical surface. However, the addition of unpaired background sweeps of different rates or direction was sufficient to alter RF characteristics across the tonotopic map in a third group of rats. These results extend earlier observations that cortical neurons can develop stimulus specific plasticity and indicate that background conditions can strongly influence cortical plasticity.
Acoustic Stimulation↗
Auditory cortex lesions and auditory-visual associative learning in cats.
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Acoustic Stimulation↗
An investigation of sensory deficits underlying the aphasia-like behavior of macaques with auditory cortex lesions.
Bilateral auditory cortex lesions in Japanese macaques result in an aphasia-like deficit in which the animals are unable to discriminate two forms of their coo vocalizations. To determine whether this deficit is sensory in nature, two monkeys with bilateral lesions were tested for their ability to discriminate frequency and frequency change. The results indicated that although the animals were able to discriminate between sounds of different frequencies, they were unable to determine whether a sound was changing in frequency. Because the animals' coo vocalizations differ primarily in the predominant direction of their frequency change and not in their absolute frequency content, the aphasia-like deficit of animals with bilateral auditory cortex lesions appears to be a sensory disorder.
Animals↗
Synaptic mechanisms of forward suppression in rat auditory cortex.
In the auditory cortex, brief sounds elicit a powerful suppression of responsiveness that can persist for hundreds of milliseconds. This forward suppression (sometimes also called forward masking) has usually been attributed to synaptic (GABAergic) inhibition. Here we have used whole-cell recordings in vivo to assess the role of synaptic inhibition in forward suppression in auditory cortex. We measured the excitatory and inhibitory synaptic conductances elicited by pairs of brief sounds presented at intervals from tens to hundreds of milliseconds. We find that inhibitory conductances rarely last longer than 50-100 ms, whereas spike responses and synaptic inputs remain suppressed for hundreds of milliseconds. We conclude that postsynaptic inhibition contributes to forward suppression for only the first 50-100 ms after a stimulus and that intracortical contributions to long-lasting suppression must involve other mechanisms, such as synaptic depression.
Acoustic Stimulation↗
Neural activity in speech-sensitive auditory cortex during silence.
That auditory hallucinations are voices heard in the absence of external stimuli implies the existence of endogenous neural activity within the auditory cortex responsible for their perception. Further, auditory hallucinations occur across a range of healthy and disease states that include reduced arousal, hypnosis, drug intoxication, delirium, and psychosis. This suggests that, even in health, the auditory cortex has a propensity to spontaneously "activate" during silence. Here we report the findings of a functional MRI study, designed to examine baseline activity in speech-sensitive auditory regions. During silence, we show that functionally defined speech-sensitive auditory cortex is characterized by intermittent episodes of significantly increased activity in a large proportion (in some cases >30%) of its volume. Bilateral increases in activity are associated with foci of spontaneous activation in the left primary and association auditory cortices and anterior cingulate cortex. We suggest that, within auditory regions, endogenous activity is modulated by anterior cingulate cortex, resulting in spontaneous activation during silence. Hence, an aspect of the brain's "default mode" resembles a (preprepared) substrate for the development of auditory hallucinations. These observations may help explain why such hallucinations are ubiquitous.
Acoustic Stimulation↗
The effect of MR scanner noise on auditory cortex activity using fMRI.
Auditory functional magnetic resonance imaging (fMRI) studies are limited by the presence of noise produced by echo planar imaging (EPI). The current study quantifies the effect of MR scanner noise on psychophysical measures of the perception of loudness and on measures of tonotopy, the representation of auditory frequencies within the auditory cortex. Seven normal hearing adults were examined using tones of 5 different auditory frequencies (250, 500, 1,000, 2,000, 4,000 Hz) of equal loudness. Using an imaging protocol with peak MR scanner noise at 1,460 Hz, the perception of loudness and detectable fMRI activity in response to a 1-kHz tone was less compared to other frequencies. When the imaging protocol was changed such that peak MR scanner noise occurred at 2,080 Hz, the perception of loudness and detectable fMRI activity in response to a 2-kHz tone was less compared to other frequencies. The reduction in the measured fMRI activity for tones near scanner frequencies may be due to an inflated scanner-induced baseline at those frequencies. In addition, fMRI activity decreased with increasing frequency, possibly due to the upward spread of masking of low-frequency, high-intensity tonal stimuli or the proximity of low-frequency core and belt areas of the auditory cortex. These results demonstrate the direct effect of scanner noise and high-intensity tonal stimuli on measurements of auditory cortex tonotopy.
Acoustic Stimulation↗
[Connection of the auditory cortex secondary zones with the auditory subcortical cerebral centers in the cat].
As it has been demonstrated microscopically, the corticofugal fibers in the AII and Ep zones of the auditory cortex in all the auditory subcortical centers (medial geniculate body, posterior colliculi of the tectum mesencephali and the superior olive nuclei) terminate by means of single axodendritic synapses, having an asymmetrically active zone, and mixed (by their form) synaptic vesicles. Small and middle dendrites make their postsynaptic part. A comparison has been carried out on distribution and form of synapses, completing the projection fibers from the zone of the primary acoustic responses (AI) and of the primary acoustic zone (AIV). Basing on the morphological data, concerning distribution and form of the synaptic terminals, a suggestion is made that physiological influence of each acoustic cortex zone is different for the medial geniculate body and posterior colliculi of the tectum mesencephali, but it is unitypical for the superior olive level.
Animals↗
Cell types in the mustached bat auditory cortex.
Cells in the auditory cortex of the mustached bat were studied with Golgi stains. No cell types appeared to be unique to the mustached bat auditory cortex or to specialized functional areas, but the laminar proportions and distributions of cell types were somewhat different from that reported for primary sensory cortex of other species. Two major cell types were distinguished, those with dendritic spines and those without spines. Non-spiny neurons were concentrated deep in layer III/IV and in layer V, and had three types of dendritic patterns, multipolar, bitufted and bipolar. Many of the non-spiny neurons were large; some nearly equaled the largest pyramidal neurons in size. Five types of spiny neurons were identified, pyramidal cells, extraverted pyramidal cells, 'spiny stellate-like' neurons, and multiform cells. In the narrow, densely packed, 'accentuated' layer II, slightly more than half of the spiny neurons were extraverted pyramidal cells, which are characterized by multiple, widely diverging apical dendrites. The high concentration of layer II extraverted pyramidal neurons is consistent with descriptions of the 'accentuated' layer II previously reported in other bat species and 'basal' insectivores. The remaining spiny neurons in layer II, and the preponderance of spiny neurons in layers III-VI, were typical pyramidal neurons that had single apical dendrites and tufts of basal dendrites. The thalamic recipient zone (deep layer III/IV) contained few candidates for spiny stellate cells, so a major constituent of the thalamic recipient zone in primary sensory cortex of many species is only a minor cellular component in the mustached bat auditory cortex.
Animals↗
Two separate codes for missing-fundamental pitch in the human auditory cortex.
Two auditory event-related potential components, the supratemporal N1 and the mismatch negativity (MMN), index traces encoding the missing-fundamental pitch. The present results suggest that these two codes derive from separate pitch extraction processes. Frequent 300-Hz and infrequent 600-Hz missing-fundamental tones were presented, in some stimulus blocks with short (150 ms), in others, with long (500 ms) stimulus durations. MMN, reflecting a preattentive change detection process, was elicited by infrequent missing-fundamental tones only in the long-duration condition. Correspondingly, subjects were able to detect these high-pitch missing-fundamental tones amongst similar low-pitch ones only when the stimulus duration was long. In addition, the MMN response peaked ca. 120 ms later for missing-fundamental tones than for pure tones of the fundamental frequency suggesting that missing-fundamental pitch resolving took substantially longer than extracting the spectral pitch. In contrast, a differential N1 response to the missing-fundamental pitch was found for both stimulus durations, with no substantial difference in peak latency between the pure and missing-fundamental tones. The contrasting features found for the two auditory cortical missing-fundamental pitch codes support the notion of two separate missing-fundamental pitch resolving mechanisms.
Adolescent↗