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Auditory processing in primate cerebral cortex.

Auditory information is relayed from the ventral nucleus of the medial geniculate complex to a core of three primary or primary-like areas of auditory cortex that are cochleotopically organized and highly responsive to pure tones. Auditory information is then distributed from the core areas to a surrounding belt of about seven areas that are less precisely cochleotopic and generally more responsive to complex stimuli than tones. Recent studies indicate that the belt areas relay to the rostral and caudal divisions of a parabelt region at a third level of processing in the cortex lateral to the belt. The parabelt and belt regions have additional inputs from dorsal and magnocellular divisions of the medial geniculate complex and other parts of the thalamus. The belt and parabelt regions appear to be concerned with integrative and associative functions involved in pattern perception and object recognition. The parabelt fields connect with regions of temporal, parietal, and frontal cortex that mediate additional auditory functions, including space perception and auditory memory.

Acoustic Stimulation↗

Determinative factors in human tonotopy: tentative analysis.

Two main parameters of stimulation were checked as an attempt to investigate the limits of the tonotopic organization in the human auditory cortex. Auditory Evoked magnetic Field (AEF) experiments from a dozen normal subjects are reported. Five of them clearly showed tonotopic AEF responses to long tone bursts stimuli, which disappeared when equivalent white noise was superimposed to the tone. Discrepancies from previous reports and high inter-individual variability are discussed.

Adolescent↗

Dynamic representational plasticity in sensory cortex.

Studies of the effects of peripheral and central lesions, perceptual learning and neurochemical modification on the sensory representations in cortex have had a dramatic effect in alerting neuroscientists and therapists to the reorganizational capacity of the adult brain. An intriguing aspect of some of these investigations, such as partial peripheral denervation, is the short-term expression of these changes. Indeed, in visual cortex, auditory cortex and somatosensory cortex loss of input from a region of the peripheral receptor epithelium (retinal, basilar and cutaneous, respectively) induces rapid expression of ectopic, or expanded, receptive fields of affected neurons and reorganization of topographic maps to fill in the representation of the denervated area. The extent of these changes can, in some cases, match the maximal extents demonstrated with chronic manipulations. The rapidity, and reversibility, of the effects rules out many possible explanations which involve synaptic plasticity and points to a capacity for representational plasticity being inherent in the circuitry of a topographic pathway. Consequently, topographic representations must be considered as manifestations of physiological interaction rather than as anatomical constructs. Interference with this interaction can produce an unmasking of previously inhibited responsiveness. Consideration of the nature of masking inhibition which is consistent with the precision and order of a topographic representation and which has a capacity for rapid plasticity requires, in addition to stimulus-driven inhibition, a source of tonic input from the periphery. Such input, acting locally to provide tonic inhibition, has been directly demonstrated in the somatosensory system and is consistent with results obtained in auditory and visual systems.

Animals↗

Augmentation of plasticity of the central auditory system by the basal forebrain and/or somatosensory cortex.

Auditory conditioning (associative learning) or focal electric stimulation of the primary auditory cortex (AC) evokes reorganization (plasticity) of the cochleotopic (frequency) map of the inferior colliculus (IC) as well as that of the AC. The reorganization results from shifts in the best frequencies (BFs) and frequency-tuning curves of single neurons. Since the importance of the cholinergic basal forebrain for cortical plasticity and the importance of the somatosensory cortex and the corticofugal auditory system for collicular and cortical plasticity have been demonstrated, Gao and Suga proposed a hypothesis that states that the AC and corticofugal system play an important role in evoking auditory collicular and cortical plasticity and that auditory and somatosensory signals from the cerebral cortex to the basal forebrain play an important role in augmenting collicular and cortical plasticity. To test their hypothesis, we studied whether the amount and the duration of plasticity of both collicular and cortical neurons evoked by electric stimulation of the AC or by acoustic stimulation were increased by electric stimulation of the basal forebrain and/or the somatosensory cortex. In adult big brown bats (Eptesicus fuscus), we made the following major findings. 1) Collicular and cortical plasticity evoked by electric stimulation of the AC is augmented by electric stimulation of the basal forebrain. The amount of augmentation is larger for cortical plasticity than for collicular plasticity. 2) Collicular and cortical plasticity evoked by AC stimulation is augmented by somatosensory cortical stimulation mimicking fear conditioning. The amount of augmentation is larger for cortical plasticity than for collicular plasticity. 3) Collicular and cortical plasticity evoked by both AC and basal forebrain stimulations is further augmented by somatosensory cortical stimulation. 4) A lesion of the basal forebrain tends to reduce collicular and cortical plasticity evoked by AC stimulation. The reduction is small and statistically insignificant for collicular plasticity but significant for cortical plasticity. 5) The lesion of the basal forebrain eliminates the augmentation of collicular and cortical plasticity that otherwise would be evoked by somatosensory cortical stimulation. 6) Collicular and cortical plasticity evoked by repetitive acoustic stimuli is augmented by basal forebrain and/or somatosensory cortical stimulation. However, the lesion of the basal forebrain eliminates the augmentation of collicular and cortical plasticity that otherwise would be evoked by somatosensory cortical stimulation. These findings support the hypothesis proposed by Gao and Suga.

Acetylcholine↗

Parallel cortical networks for volitional control of swallowing in humans.

A number of studies have demonstrated the involvement of parallel networks in the control of voluntary sequential motor procedures. We sought to determine whether a parallel network organization may be found for complex, sequentially based motor systems that are the product of both voluntary and automatic control processes. Specifically, we sought to determine whether the cortical organizational scheme for voluntary repetitive swallowing in adult humans is characterized by a hierarchical dual-projection model or by modules organized into parallel systems. We utilized functional magnetic resonance imaging (fMRI) to investigate cortical function during normal swallowing tasks in eight healthy human adults. Subjects performed both dry (saliva) and bolus (3 ml/bolus of water) swallows. Activation during swallowing tasks localized to sensorimotor areas (M1, S1, and SMA), S2, premotor cortex, posterior parietal cortex, cingulate gyrus, inferior frontal gyrus, the cerebellum, the insular cortex, auditory cortex, corpus callosum, and the basal ganglia and thalamus. Principal components analysis (PCA) of these regions revealed five functional clusters or modules: (1) sensorimotor areas and cingulate gyrus; (2) inferior frontal gyrus, S2, corpus callosum, basal ganglia and thalamus; (3) premotor cortex and posterior parietal cortex; (4) cerebellum; and (5) insula. Analysis of the functional relationship between these areas demonstrated two parallel loops defined by connections to either the cerebellum or insula and connected through the sensorimotor-cingulate module. Path analysis was performed to test the hypothesis of modules organized into parallel loops versus a hierarchical dual-projection model consisting of two separate, singular hierarchical serial pathways from the sensorimotor cortex or insula to the thalamus. These results support the model of modules organized into parallel loops (P=0.8), but not the hierarchical dual-projection model (P<0.0001). Organization of the control of voluntary repetitive swallowing into two parallel systems may confer the ability to effectively coordinate and integrate this highly complex sequentially based motor behavior.

Adult↗

Cortical and subcortical EEG patterns during moderate-intensity barbiturate withdrawal.

Electroencephalograms were recorded from cortical and subcortical brain regions during moderate-intensity barbiturate withdrawal in the cat. Physical dependence was produced by administration of Na-barbital by the "low dose barbiturate" treatment schedule. During moderate-intensity withdrawal, the EEG pattern showed 4- to 6-Hz rhythmic bursts in the hippocampus, visual cortex, auditory cortex, motor cortex, nucleus centralis centralis of the thalamus, caudate nucleus, and lateral geniculate body. The bursts in the hippocampus were spike-and-wave type. In animals which had spontaneous seizures, 4- to 6-Hz bursts occurred in the motor cortex, whereas this abnormality did not occur in the motor cortex in animals which did not have spontaneous generalized seizures. In addition, prolonged high-frequency, high-voltage discharges and spikes were recorded from the amygdala during withdrawal. Spontaneous generalized or partial seizures occurred during withdrawal; the abnormal EEG patterns associated with seizures appeared to originate in various subcortical sites. Perhaps the most important result was the finding that no one brain region or set of brain regions seemed to be specifically involved in barbiturate withdrawal.

Animals↗

Spatio-temporal source modeling of evoked potentials to acoustic and cochlear implant stimulation.

Spatio-temporal source modeling (STSM) of event-related potentials was used to estimate the loci and characteristics of cortical activity evoked by acoustic stimulation in normal hearing subjects and by electrical stimulation in cochlear implant (CI) subjects. In both groups of subjects, source solutions obtained for the N1/P2 complex were located in the superior half of the temporal lobe in the head model. Results indicate that it may be possible to determine whether stimulation of different implant channels activates different regions of cochleotopically organized auditory cortex. Auditory system activation can be assessed further by examining the characteristics of the source wave forms. For example, subjects whose cochlear implants provided auditory sensations and normal hearing subjects had similar source activity. In contrast, a subject in whom implant activation evoked eyelid movements exhibited different source wave forms. STSM analysis may provide an electrophysiological technique for guiding rehabilitation programs based on the capabilities of the individual implant user and for disentangling the complex response patterns to electrical stimulation of the brain.

Acoustic Stimulation↗

Transient induction of neuronal nitric oxide synthase in neurons of rat cerebral cortex after status epilepticus.

The change in neuronal nitric oxide synthase (nNOS) expression after status epilepticus induced by kainate was examined in the rat cerebral cortex. Expressional change was assessed using nNOS immunohistochemistry and reverse transcription-polymerase chain reaction for nNOS mRNA. Constitutive nNOS-positive neurons was observed in the cerebral cortex of the control group. At 1 and 3 days after status epilepticus, nNOS-positive neurons were present in a deep layer of various cortical regions such as primary motor cortex, secondary motor cortex, primary somatosensory cortex, secondary somatosensory cortex, parietal association cortex, insular cortex, ectorhinal cortex, temporal association cortex, auditory cortex and visual cortex. The level of nNOS mRNA increased at 1, 3, 6 and 12 days after status epilepticus compared to controls. This report provides the first morphological evidence that nNOS are induced in neurons of the cerebral cortex following seizure.

Animals↗

[Measurement of noise protection in functional magnetic resonance imaging].

BACKGROUND: Functional magnetic resonance imaging (fMRI) can detect changes in oxygen saturation of the brain. Fast changing high gradient fields are necessary which produce high levels of noise. In studies of the auditory cortex, auditory stimuli have to be perceived and discriminated against the noise level of the activated tomograph. MATERIAL AND METHODS: The generated frequency bands and their intensities during fMRI with a Siemens Magnetom Vision, 1.5 T, EPI sequence were measured in the outer ear canal of a dummy head. Noise attenuation was evaluated with four different noise muffs (simple/inexpensive products, quality product, specialized fMRI muffs). RESULTS: Without protection, peak noise levels reached up to 111 dB(A) near 1000 Hz in the dummy ear canal. Major noise attenuation was only found at higher frequencies (4000 Hz by about 25 dB; 8000 Hz by about 35 dB) with the quality product and the specialized fMRI muffs. CONCLUSION: Only quality noise products can sufficiently protect patients from high sound pressure levels of tomograph noise. If in the future higher gradient fields are applied at faster slew rates, acoustic stimuli can safely be applied only in combination with increased hearing protection systems in order to minimize the risk of noise trauma.

Ear Protective Devices↗

Diurnal variation of cation pump enzyme activity in pineal and seven other rat brain regions.

Adult female Long-Evans rats were maintained on an automatically regulated artificial lighting schedule of light:dark (L:D) 14.5:9.5 for 12 wk. After sacrifice at 0630, 1130, 1600, 1800, 2000, 2200, 0230, or 0400, the pineals were removed, weighed, and assayed for N-acetyltransferase (NAT), melatonin, Mg++-paranitrophenylphosphatase (pNPPase), and K-pNPPase activity. The brains were quickly dissected into the following areas: cerebellum, superior colliculi, inferior colliculi, visual cortex, auditory cortex, sensorimotor cortex, and the hypothalamic area around the suprachiasmatic nucleus. These regions were weighed and 10% sucrose homogenates were prepared for determinations of protein, Mg++-pNPPase, and K+-pNPPase activity. Pineal melatonin rose over six-fold from 144 +/- 70 pg/gland at 1130 to 981 +/- 173 pg/gland at 0230. Similarly, pineal NAT activity rose over 11-fold, from 119 +/- 12 pmol/gland/h to 1315 +/- 232 pmol/gland/h at the same times. K+-pNPPase activity rose by about two-thirds, from 133 +/- 12.8 nmol/gland/h to 224 +/- 22.3 nmol/gland/h from 1600 to 0230. However, when expressed per mg protein, these differences in pNPPase activity were not significant. There were no significant daily rhythms discernible in any of the seven other brain regions across these times. We conclude that cation pump enzyme activity varies only slightly with time in the rat brain and pineal gland, in spite of definite daily rhythms of pineal melatonin and NAT activity.

4-Nitrophenylphosphatase↗

The spatiotemporal pattern of auditory cortical responses during verbal hallucinations.

Functional magnetic resonance imaging (fMRI) studies can provide insight into the neural correlates of hallucinations. Commonly, such studies require self-reports about the timing of the hallucination events. While many studies have found activity in higher-order sensory cortical areas, only a few have demonstrated activity of the primary auditory cortex during auditory verbal hallucinations. In this case, using self-reports as a model of brain activity may not be sensitive enough to capture all neurophysiological signals related to hallucinations. We used spatial independent component analysis (sICA) to extract the activity patterns associated with auditory verbal hallucinations in six schizophrenia patients. SICA decomposes the functional data set into a set of spatial maps without the use of any input function. The resulting activity patterns from auditory and sensorimotor components were further analyzed in a single-subject fashion using a visualization tool that allows for easy inspection of the variability of regional brain responses. We found bilateral auditory cortex activity, including Heschl's gyrus, during hallucinations of one patient, and unilateral auditory cortex activity in two more patients. The associated time courses showed a large variability in the shape, amplitude, and time of onset relative to the self-reports. However, the average of the time courses during hallucinations showed a clear association with this clinical phenomenon. We suggest that detection of this activity may be facilitated by examining hallucination epochs of sufficient length, in combination with a data-driven approach.

Acoustic Stimulation↗

Requirement of the auditory association cortex for discrimination of vowel-like sounds in rats.

We investigated the roles of the auditory cortex in discrimination learning of vowel-like sounds consisting of multiple formants. Rats were trained to discriminate between synthetic sounds with four formants. Bilateral electrolytic lesions including the primary auditory cortex and the dorsal auditory association cortex impaired multiformant discrimination, whereas they did not significantly affect discrimination between sounds with a single formant or between pure tones. Local lesions restricted to the dorsal/rostral auditory association cortex were sufficient to attenuate multiformant discrimination learning, and lesions restricted to the primary auditory cortex had no significant effects. These findings indicate that the dorsal/rostral auditory association cortex but not the primary auditory cortex is required for discrimination learning of vowel-like sounds with multiple formants in rats.

Acoustic Stimulation↗

Identification of a pathway for intelligible speech in the left temporal lobe.

It has been proposed that the identification of sounds, including species-specific vocalizations, by primates depends on anterior projections from the primary auditory cortex, an auditory pathway analogous to the ventral route proposed for the visual identification of objects. We have identified a similar route in the human for understanding intelligible speech. Using PET imaging to identify separable neural subsystems within the human auditory cortex, we used a variety of speech and speech-like stimuli with equivalent acoustic complexity but varying intelligibility. We have demonstrated that the left superior temporal sulcus responds to the presence of phonetic information, but its anterior part only responds if the stimulus is also intelligible. This novel observation demonstrates a left anterior temporal pathway for speech comprehension.

Acoustic Stimulation↗

Representation of spectral and temporal sound features in three cortical fields of the cat. Similarities outweigh differences.

This study investigates the degree of similarity of three different auditory cortical areas with respect to the coding of periodic stimuli. Simultaneous single- and multiunit recordings in response to periodic stimuli were made from primary auditory cortex (AI), anterior auditory field (AAF), and secondary auditory cortex (AII) in the cat to addresses the following questions: is there, within each cortical area, a difference in the temporal coding of periodic click trains, amplitude-modulated (AM) noise bursts, and AM tone bursts? Is there a difference in this coding between the three cortical fields? Is the coding based on the temporal modulation transfer function (tMTF) and on the all-order interspike-interval (ISI) histogram the same? Is the perceptual distinction between rhythm and roughness for AM stimuli related to a temporal versus spatial representation of AM frequency in auditory cortex? Are interarea differences in temporal response properties related to differences in frequency tuning? The results showed that: 1) AM stimuli produce much higher best modulation frequencies (BMFs) and limiting rates than periodic click trains. 2) For periodic click trains and AM noise, the BMFs and limiting rates were not significantly different for the three areas. However, for AM tones the BMF and limiting rates were about a factor 2 lower in AAF compared with the other areas. 3) The representation of stimulus periodicity in ISIs resulted in significantly lower mean BMFs and limiting rates compared with those estimated from the tMTFs. The difference was relatively small for periodic click trains but quite large for both AM stimuli, especially in AI and AII. 4) Modulation frequencies <20 Hz were represented in the ISIs, suggesting that rhythm is coded in auditory cortex in temporal fashion. 5) In general only a modest interdependence of spectral- and temporal-response properties in AI and AII was found. The BMFs were correlated positively with characteristic frequency in AAF. The limiting rate was positively correlated with the frequency-tuning curve bandwidth in AI and AII but not in AAF. Only in AAF was a correlation between BMF and minimum latency was found. Thus whereas differences were found in the frequency-tuning curve bandwidth and minimum response latencies among the three areas, the coding of periodic stimuli in these areas was fairly similar with the exception of the very poor representation of AM tones in AII. This suggests a strong parallel processing organization in auditory cortex.

Acoustic Stimulation↗

Auditory-somatosensory multisensory processing in auditory association cortex: an fMRI study.

Using high-field (3 Tesla) functional magnetic resonance imaging (fMRI), we demonstrate that auditory and somatosensory inputs converge in a subregion of human auditory cortex along the superior temporal gyrus. Further, simultaneous stimulation in both sensory modalities resulted in activity exceeding that predicted by summing the responses to the unisensory inputs, thereby showing multisensory integration in this convergence region. Recently, intracranial recordings in macaque monkeys have shown similar auditory-somatosensory convergence in a subregion of auditory cortex directly caudomedial to primary auditory cortex (area CM). The multisensory region identified in the present investigation may be the human homologue of CM. Our finding of auditory-somatosensory convergence in early auditory cortices contributes to mounting evidence for multisensory integration early in the cortical processing hierarchy, in brain regions that were previously assumed to be unisensory.

Acoustic Stimulation↗

Primary auditory pathway and reticular activating system dysfunction in Alzheimer's disease.

Patients with Alzheimer's disease (AD) have pathologic involvement of several important components of the primary auditory pathway, including the inferior colliculus, medial geniculate body, primary auditory cortex, and secondary auditory cortex. The main components of the brainstem auditory evoked response (BAER) and middle latency response (MLR) reflect the function of portions of the primary auditory pathway, including those affected pathologically in AD. The amplitude of the P1 component of the MLR reflects the degree of neuronal activity of midbrain portions of the ascending reticular activating system (ARAS) with cortical cholinergic projections. To determine whether there is dysfunction of the primary auditory pathway and ARAS in AD, we compared simultaneous BAER and MLR component latency and amplitude measurements in patients with mild-moderate AD (n = 35) and age-matched healthy elderly controls (n = 34). There were significant latency delays in brainstem transmission time (BAER I-V interpeak latency; p < 0.05) and in primary auditory cortex evoked potential generation (MLR Pa latency; p < 0.05) in the AD group compared with controls. In addition, there was a significant reduction in the P1 component amplitude of the MLR in the AD group (p < 0.01). These results indicate dysfunction of the primary auditory pathway and ARAS in patients with mild-moderate AD and support the hypothesis that impairment of auditory function and of arousal are intrinsic features of AD.

Aged↗