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Plasticity of auditory cortex associated with sensorineural hearing loss in adult C57BL/6J mice.

The representation of frequency was mapped in the primary auditory cortex (AI) of C57BL/6J (C57) mice during young adulthood (1.5-2 months) when hearing is optimal, and at 3, 6, and 12 months of age, a period during which progressive, high frequency, sensorineural hearing loss occurs in this strain. Maps were also obtained from CBA/CaJ mice which retain good hearing as they age. In AI of young adult C57 mice and CBA mice, characteristic frequencies (CFs) of multiple-unit clusters were easily identified with extracellular recordings, and a general tonotopic organization was observed from dorsal (high frequency) to ventral and caudal (low frequency). In individual cases there appeared to be deviations from the above tonotopic organization, despite the fact that inbred mice are genetically invariant. As progressive loss of high frequency sensitivity ensued peripherally, a substantially increased representation of middle frequencies was observed in AI. There was no apparent change in the surface area of the auditory cortex despite the elimination of high frequencies, and virtually the entire auditory cortex became devoted to the middle frequencies (especially 10-13 kHz) for which sensitivity remained high. Similar age-related changes were not observed in normal-hearing CBA mice. These findings indicate that plasticity in the representation of frequency in AI is associated with high frequency hearing loss in C57 mice.

Acoustic Stimulation↗

Projections from the auditory cortex to the superior olivary complex in guinea pigs.

We used anterograde tracing techniques to characterize projections from auditory cortex to the superior olivary complex (SOC) in guinea pigs. Large injections of fluorescent or biotinylated dextrans into the temporal cortex labeled many axons in the SOC. Labeled boutons were most numerous in the ventral nucleus of the trapezoid body, with additional boutons in all other olivary nuclei. The distribution of boutons was similar in the ipsilateral and contralateral SOC; however, the contralateral SOC had markedly fewer axons and boutons. Similar patterns of labeling were also observed following injections confined to primary auditory cortex or the dorsocaudal auditory field. Cortical axons in many of the SOC nuclei share numerous morphological features, suggesting that individual axons may innervate multiple nuclei and have widespread effects. In addition, some nuclei contain axons with branching or termination patterns unique to that nucleus; these axons may represent focused projections with effects limited to individual SOC nuclei. Given the many projections of SOC nuclei, cortico-olivary projections are in a position to modify the activity of many brainstem auditory circuits.

Animals↗

Effects of musical training on the auditory cortex in children.

Several studies of the effects of musical experience on sound representations in the auditory cortex are reviewed. Auditory evoked potentials are compared in response to pure tones, violin tones, and piano tones in adult musicians versus nonmusicians as well as in 4- to 5-year-old children who have either had or not had extensive musical experience. In addition, the effects of auditory frequency discrimination training in adult nonmusicians on auditory evoked potentials are examined. It was found that the P2-evoked response is larger in both adult and child musicians than in nonmusicians and that auditory training enhances this component in nonmusician adults. The results suggest that the P2 is particularly neuroplastic and that the effects of musical experience can be seen early in development. They also suggest that although the effects of musical training on cortical representations may be greater if training begins in childhood, the adult brain is also open to change. These results are discussed with respect to potential benefits of early musical training as well as potential benefits of musical experience in aging.

Auditory Cortex↗

The characteristics of acetylcholine release mechanisms in the auditory cortex.

1. The characteristics of the acetylcholine (ACh) release mechanism have been studied in the auditory cortex of rabbits on stimulation of the ipsilateral medial geniculate nucleus.2. On stimulation of the medial geniculate nucleus the mean release of ACh from the auditory receiving cortex was 6.1 times the spontaneous release; the mean release from other parts of the cortex was 2.2 times the spontaneous release.3. The frequency of stimulation most effective in evoking ACh release was found to be 10/sec.4. Both the spontaneous and evoked release of ACh were reduced by 40-65% in the absence of calcium from the solution bathing the auditory cortex, and increased by 15-25% when the calcium concentration in the bathing solution was doubled.5. The presence of low concentrations of magnesium in the fluid bathing the cortex was essential for the optimal release of ACh, but high magnesium concentrations lowered this release.6. The presence of triethylcholine (TEC) in the fluid bathing the auditory cortex reduced both the spontaneous and evoked release of ACh. This reduction was reversed in the presence of choline.7. The effects of calcium, magnesium and TEC on the ACh release mechanism in the cerebral cortex and at the neuromuscular junction are compared.

Acetylcholine↗

Separate neuronal populations of the rat globus pallidus projecting to the subthalamic nucleus, auditory cortex and pedunculopontine tegmental area.

The topographic arrangement of globus pallidus neurons sending axons to the subthalamic nucleus, auditory cortex and pedunculopontine tegmental nucleus was studied in the rat using retrograde fluorescent tracers. Neurons projecting to the subthalamic nucleus were localized in the rostral part of the globus pallidus, while neurons projecting to the auditory cortex and to the pedunculopontine tegmental nucleus were located in the caudal part. The two populations of pallidocortical and pallidotegmental neurons were also distributed in a separate manner within the caudal globus pallidus. The former neurons were large and located more ventromedially, whereas the latter were medium-sized and located more dorsolaterally. Using a retrograde fluorescent tracing technique combined with choline acetyltransferase immunofluorescence histochemistry, it was found that a vast majority of pallidocortical neurons expressed choline acetyltransferase immunoreactivity, and that pallidotegmental neurons rarely exhibited choline acetyltransferase immunoreactivity. A method of retrograde tracing with wheatgerm agglutinin conjugated with horseradish peroxidase associated to immunohistochemistry for glutamate decarboxylase confirmed the GABAergic nature of the pallidotegmental pathway. The present study revealed the independent nature of the globus pallidus neurons projecting to the subthalamic nucleus, auditory cortex and pedunculopontine tegmental nucleus. Within this cellular arrangement, the presence of functionally distinct neuronal populations at the caudal pallidal level was also identified, with large cholinergic cells innervating the neocortex and medium-sized GABAergic cells "feeding" the mesencephalic tegmentum.

Animals↗

Response linearity in primary auditory cortex of the ferret.

The responses of neurons within the primary auditory cortex (A1) of the ferret elicited by broadband dynamic spectral ripple stimuli were examined over a range of ripple spectral densities and ripple velocities. The large majority of neurons showed modulated responses to these stimuli and responded most strongly at low ripple densities and velocities. The period histograms of their responses were subjected to Fourier analysis, and the ratio of the magnitudes of the f1 and fo (DC) components of these responses were calculated to give a quantitative index of response linearity. For 82 out of 396 neurons tested (20.7%) this ratio remained above 1.0 over the entire range of ripple densities and velocities. These neurons were classified as 'consistently linear'. A further 134/396 (33.8%) of neurons maintained an f1/f0 ratio above 1.0 for either a range of ripple densities at a fixed ripple velocity, or over a range of ripple velocities at a specific ripple density, and were classified as 'locally linear'. Interestingly, for the superficial layers of the primary auditory cortex, consistently linear and locally linear neurons outnumbered nonlinear neurons by a 2:1 ratio. The converse was true for the deep layers. Unlike in primary visual cortex, where f1/f0 ratios have been reported to exhibit a bimodal distribution with a minimum at f1/f0 = 1, f1/f0 ratios for A1 are unimodally distributed with a peak at f1/f0 = 1.

Acoustic Stimulation↗

[Responses of neurons in isolated strips of cat auditory cortex to intracortical stimulation].

Neuron responses in an isolated slab of the auditory cortex (A1 zone) to intracortical electric stimulation of its layer IV have been investigated extracellularly in acute experiments on cats immobilized with d-tubocurarin. Responding neurons have been found in all layers of the slab. Their distribution in the slab depth depended on the distances between the recording and stimulating electrodes. The latent periods of responses of different neurons varied between 0.8 and 25 ms. Most of neurons were involved in the reaction mono- or disynaptically within the employed limits of distances between the electrodes (0.5-2 mm). However, many neurons responded polysynaptically. This indicates a complex character of interneuron interactions even in a limited section of the brain cortex. No after-discharges with latent periods above 40 ms have been recorded in the isolated auditory cortex slab in response to intracortical stimulation.

Animals↗

Representation of the two ears in the auditory cortex: a re-examination.

The current theory of the representation of the two ears in the auditory cortex, namely that both ears are represented in each cerebral cortex and that the contralateral ear receives a stronger representation, is critically reviewed in the light of more recent evidence. On the basis of neurophysiological and behavioral studies in animals and a variety of clinical studies, it is concluded that the notion of contralateral ear dominance in the cortex requires revision, and it is proposed that the contralateral sound field, rather than contralateral ear as such, is represented in each cortex. It is suggested that the contralateral and ipsilateral inputs to the auditory cortex do not so much differ in their relative strengths, but in the qualitative nature of the information they transmit.

Animals↗

Multisensory integration of dynamic faces and voices in rhesus monkey auditory cortex.

In the social world, multiple sensory channels are used concurrently to facilitate communication. Among human and nonhuman primates, faces and voices are the primary means of transmitting social signals (Adolphs, 2003; Ghazanfar and Santos, 2004). Primates recognize the correspondence between species-specific facial and vocal expressions (Massaro, 1998; Ghazanfar and Logothetis, 2003; Izumi and Kojima, 2004), and these visual and auditory channels can be integrated into unified percepts to enhance detection and discrimination. Where and how such communication signals are integrated at the neural level are poorly understood. In particular, it is unclear what role "unimodal" sensory areas, such as the auditory cortex, may play. We recorded local field potential activity, the signal that best correlates with human imaging and event-related potential signals, in both the core and lateral belt regions of the auditory cortex in awake behaving rhesus monkeys while they viewed vocalizing conspecifics. We demonstrate unequivocally that the primate auditory cortex integrates facial and vocal signals through enhancement and suppression of field potentials in both the core and lateral belt regions. The majority of these multisensory responses were specific to face/voice integration, and the lateral belt region shows a greater frequency of multisensory integration than the core region. These multisensory processes in the auditory cortex likely occur via reciprocal interactions with the superior temporal sulcus.

Acoustic Stimulation↗

Functional organization of the callosal connections of the cat auditory cortex.

In acute experiments on immobilized cats, using a method of topographical recording of homotopic and heterotopic transcallosal responses, the functional organization of the callosal connections of the auditory cortex was investigated. It was established that the homotopic potentials of the primary projection field (AI) have the greatest amplitude, minimal temporal parameters, and the maximal stability of these characteristics as compared with the associative fields of the auditory cortex (AII, AIV, Ep). The heterotropic transcallosal responses in field AI appeared during stimulation of the analogous field, while in field Ep, they were recorded both during stimulation of the analogous field, and of fields AI and AII of the opposite hemisphere. It is hypothesized that the structure of the transcallosal connections of the primary projection field s of the auditory cortex is characterizised by homotopy, whereas in the associative auditory fields the role of heterotopic transcallosal interactions increases. It is possible that such a structure of the transcallosal connections assures a significant role for interhemispheric interactins in the mechanisms of spatial audition.

Animals↗

[The structural-functional organization of the auditory cortex in rats].

Using axonal transport of horseradish peroxidase and electrophysiological mapping technique, studies have been made on structural and functional organization of the auditory cortex in rats. After the injection of HRP to peripheral parts of the auditory cortex, mainly initial neurones were found in the dorsal and median parts of the geniculate body. Electrophysiological experiments revealed the localization of neurones with widespread frequency-threshold curves and high thresholds at characteristic frequency. On the basis of the data obtained, it is suggested that similar to other mammals, rats have the areas of secondary fields of the auditory cortex which surround central coniocortex.

Animals↗

No indication of brain reorganization after unilateral ischemic lesions of the auditory cortex.

We used magnetoencephalography to study contralesional auditory reorganization in three men with chronic unilateral ischemic lesions of the auditory cortex. Although no response was found over the lesioned hemisphere, processing in the unaffected hemisphere was indistinguishable vs healthy controls. In contrast to sensorimotor and language systems, the auditory system seems to lack contralateral reorganization, presumably because patients are typically not aware of hearing deficits and thus do not perform training.

Acoustic Stimulation↗

Spatial processing in the auditory cortex of the macaque monkey.

The patterns of cortico-cortical and cortico-thalamic connections of auditory cortical areas in the rhesus monkey have led to the hypothesis that acoustic information is processed in series and in parallel in the primate auditory cortex. Recent physiological experiments in the behaving monkey indicate that the response properties of neurons in different cortical areas are both functionally distinct from each other, which is indicative of parallel processing, and functionally similar to each other, which is indicative of serial processing. Thus, auditory cortical processing may be similar to the serial and parallel "what" and "where" processing by the primate visual cortex. If "where" information is serially processed in the primate auditory cortex, neurons in cortical areas along this pathway should have progressively better spatial tuning properties. This prediction is supported by recent experiments that have shown that neurons in the caudomedial field have better spatial tuning properties than neurons in the primary auditory cortex. Neurons in the caudomedial field are also better than primary auditory cortex neurons at predicting the sound localization ability across different stimulus frequencies and bandwidths in both azimuth and elevation. These data support the hypothesis that the primate auditory cortex processes acoustic information in a serial and parallel manner and suggest that this may be a general cortical mechanism for sensory perception.

Acoustic Stimulation↗

Long-term potentiation in the auditory cortex of adult rats.

Long-term potentiation (LTP) in the auditory cortex was studied in slices obtained from adult rats. White matter stimulation produced field potentials in layers II/III, which were composed of two negative waves followed by a slow positivity. The second negative and third positive waves were blocked by a low Ca2+ (0.48 mM) medium or by 6-cyano-7-nitroquinoxaline-2,3-dione (CNQX, 10 microM), while the first negativity unchanged. D-2-amino-5-phosphonovalerate (D-AP5, 50 microM) showed no clear effect on the field potentials. Tetanic stimulation of the white matter produced potentiation in the second negativity, and the potentiation was maintained for at least 30 min. D-AP5 blocked this potentiation completely. These data indicate that LTP is evoked by activation of N-methyl-D-aspartate (NMDA) receptors in the auditory cortex of adult rats.

2-Amino-5-phosphonovalerate↗

Neonatal auditory cortex lesions result in aberrant crossed corticotectal and corticothalamic projections in rats.

Ablation of one auditory cortex at birth in rats results in the formation of aberrant crossed projections from the intact hemisphere. These aberrant projections extend throughout most of the corticorecipient zone of the contralateral inferior colliculus and can be traced as far rostral as the contralateral medial geniculate nucleus. The aberrant crossed corticothalamic axons arise from layer V pyramidal neurons in the intact auditory cortex.

Animals↗

Neural connectivity only accounts for a small part of neural correlation in auditory cortex.

In order to allow the relation of functional connectivity patterns (inferred from cross-correlograms) to structural connectivity (the anatomical substrate), we analyzed cross-correlogram peaks for spontaneous and stimulated activity in the auditory cortex. It was assumed that the broad correlograms, usually encountered, represent neural connectivity as well as secondary effects such as intrinsic firing patterns, global synchrony related to the ongoing electroencephalographic activity, and stimulus-related effects. Data were collected from 604 neuron pairs recorded under spontaneous conditions in primary auditory cortex of seven juvenile (30-70 days) and nine adult cats. Three hundred and six pairs (51%) had a peak cross-correlation coefficient significantly different from zero. For 113 neuron pairs out of this subgroup, correlations were calculated also for spike trains recorded during click stimulation. After a combined burst-correction and deconvolution procedure was carried out, the correlation peak strengths were not significantly changed for spontaneous activity, but peak width was narrower for single-electrode pairs than for dual-electrode pairs, suggesting a better synchronization for neighboring neurons. Under click stimulation conditions, overall peak synchronization strength was independent of interelectrode distance, whereas, after correction for secondary and stimulus effects, peak synchronization was significantly lower for dual-electrode pairs. However, the primary peak width for single-electrode pairs under stimulus conditions was no longer different from that of dual-electrode pairs. This implies that both under spontaneous and stimulus conditions secondary effects largely obscure any underlying correlation produced by anatomical connectivity. The secondary effects may be the result of intrinsic as well as network properties in auditory cortex and may functionally be more important than the weak primary effects resulting from anatomical connections. Cross-interval analysis suggests that the correlations in auditory cortex are dynamic and may show random switching between states of stronger and weaker synchronization.

Acoustic Stimulation↗

Responses of neurons to click-pairs as simulated echoes: auditory nerve to auditory cortex.

When two identical sounds are presented from different locations with a short interval between them, the perception is of a single sound source at the location of the leading sound. This "precedence effect" is an important behavioral phenomenon whose neural basis is being increasingly studied. For this report, neural responses were recorded to paired clicks with varying interstimulus intervals, from several structures of the ascending auditory system in unanesthetized animals. The structures tested were the auditory nerve, anteroventral cochlear nucleus, superior olivary complex, inferior colliculus, and primary auditory cortex. The main finding is a progressive increase in the duration of the suppressive effect of the leading sound (the conditioner) on the response to the lagging sound (the probe). The first major increase occurred between the lower brainstem and inferior colliculus, and the second between the inferior colliculus and auditory cortex. In neurons from the auditory nerve, cochlear nucleus, and superior olivary complex, 50% recovery of the response to the probe occurred, on average, for conditioner and probe intervals of approximately 2 ms. In the inferior colliculus, 50% recovery occurred at an average separation of approximately 7 ms, and in the auditory cortex at approximately 20 ms. Despite these increases in average recovery times, some neurons in every structure showed large responses to the probe within the time window for precedence (approximately 1-4 ms for clicks). This indicates that during the period of the precedence effect, some information about echoes is retained. At the other extreme, for some cortical neurons the conditioner suppressed the probe response for intervals of up to 300 ms. This is in accord with behavioral results that show dominance of the leading sound for an extended period beyond that of the precedence effect. Other transformations as information ascended included an increased variety in the shapes of the recovery functions in structures subsequent to the nerve, and neurons "tuned" to particular conditioner-probe intervals in the auditory cortex. These latter are reminiscent of neurons tuned to echo delay in bats, and may contribute to the perception of the size of the acoustic space.

Acoustic Stimulation↗

Enhancement of the auditory cortex evoked responses in awake guinea pigs after noise exposure.

In a previous paper [Popelár et al., Hear. Res. 26, 239-247 (1987)] we have shown that amplitudes of the auditory cortex evoked responses (AC-ER) in awake guinea pigs were enhanced for several hours after 1 h of noise exposure whereas amplitudes of the compound potential of the auditory nerve (CAP) and of the inferior colliculus evoked responses (IC-ER) declined. The present study demonstrates that the duration of the AC-ER amplitude increase is related to the intensity of the noise exposure (white noise, for 30 min or 1 h, intensity range 105-125 dB). The AC-ER amplitude as well as the threshold shift increased linearly with increasing intensity of the noise. The maximum AC-ER increase occurred when clicks served as stimuli; amplitude enhancement was smaller for 1 kHz tone pips and was absent when 20 kHz tone pips were used. The amplitude enhancement was specific for the auditory cortex since the amplitude of visually evoked responses, recorded in the occipital cortex, was unchanged after noise exposure. It is suggested that the postexposure amplitude enhancement of the AC-ER is produced by temporary exhaustion of inhibitory processes in the auditory cortex.

Acoustic Stimulation↗