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Effect of tone-pulse rise time on rate-level functions of cat auditory cortex neurons: excitatory and inhibitory processes shaping responses to tone onset.

1. The responses of cat auditory cortex neurons are largely dominated by transient stimulus events, including tone-pulse onset. In addition, these neurons often receive sensitive inhibitory inputs in tone frequency-intensity domains flanking the excitatory one centered at characteristic frequency (CF). These observations suggest that auditory cortex neurons might be sensitive to the spectral splatter that occurs at tone onset due to the tone-pulse envelope shape. 2. To investigate this hypothesis, single neurons in the primary auditory cortex of anesthetized cats were studied for the form of their spike-rate versus tone-level functions using CF tone pulses of different rise times. Stimuli were presented to the contralateral ear using a calibrated, sealed stimulus delivery system. 3. Some neurons with monotonic rate-level functions for conventional (5-10 ms) rise-time tones were relatively insensitive to variations in tone-pulse rise time. Other monotonic neurons showed rate-level functions that became increasingly bell shaped for shorter rise-time stimuli. All neurons with bell-shaped, nonmonotonic rate-level functions for conventional rise-time tones became increasingly nonmonotonic for shorter rise-time signals. In the same neurons, lengthening of tone rise times typically reduced the slope of the high-intensity, descending limb of the rate-level function, in some cases to zero. 4. This pattern of rise-time effects is consistent with previous evidence on the association between rate-level function shape and the presence of inhibitory tone response areas flanking the excitatory one at CF. The present data suggest that cortical neurons are sensitive to the gross shape of the short-term stimulus spectrum at tone onset, and that for many neurons, the nonmonotonic form of CF tone rate level functions may be configured as much by the rate of tone onset as by the plateau amplitude of a tone pulse.

Acoustic Stimulation

Cortical connections of electrophysiologically and architectonically defined subdivisions of auditory cortex in squirrels.

Multiunit recordings with microelectrodes were used to identify and delimit subdivision of auditory cortex in squirrels. In the same animals, cortical connections of subdivisions of auditory cortex were determined by placing injections of the tracer wheat germ agglutinin conjugated with horseradish peroxidase (WGA-HRP) into electrophysiologically defined locations. The electrophysiological results and patterns of connections were later related to myeloarchitectonic distinctions in brain sections cut parallel to the surface of the artificially flattened cortex. As previously described (Merzenich et al.: J. Comp. Neurol. 166:387-402, '76), a primary auditory field, A-I, was characterized by (1) neurons narrowly tuned to tone frequency; (2) a tonotopic map with high frequencies, which represented caudal to low frequencies; and (3) dense myelination. A-I was reciprocally connected with a rostral field, R, a parietal ventral somatosensory representation, PV, cortex ventral to A-I, and other nearby regions of cortex of the same hemisphere. Callosal connections of A-I were with A-I, R, and two or more other regions of temporal cortex. The less densely myelinated rostral field, R, also had neurons that were frequency tuned, but the neurons were often less securely driven. R appeared to have a tonotopic organization that roughly mirrored that of A-I. Ipsilateral connections of R included A-I, PV, and cortex ventral and caudal to R. Callosal connections were with R, A-I, PV, and cortex ventral and caudal to R. Callosal connections were with R, A-I, PV, and other locations in temporal cortex. Cortex in caudal PV, ventral to A-I, and ventral to R was responsive to auditory stimuli, but responses to pure tones were weak and inconsistent, and habituation to a repeated stimulus was rapid. The cortex responsive to auditory stimuli included some but not all of the cortex connected with A-I and R. The results lead to the conclusion that auditory cortex of squirrels contains at least two tonotopically organized fields, possibly as many as five or more auditory fields, and at least two auditory-somatosensory fields.

Animals

Primary auditory cortex in the rat: transient expression of acetylcholinesterase activity in developing geniculocortical projections.

A characteristic pattern of acetylcholinesterase (AChE) activity is expressed transiently in primary auditory cortex (cortical area 41) of developing laboratory rats during early postnatal life. This AChE activity occurs as a dense plexus in cortical layer IV and the deep part of layer III. This transient band of AChE activity is first detected by histochemical techniques on postnatal day (P) 3, reaches peak intensity at approximately P8-10, and declines to form the adult pattern by P23. The ventral nucleus of the medial geniculate body of the thalamus also displays prominent, and transient, staining for AChE. This intense staining for AChE, found within neuronal somata and neuropil, is detected at the time of birth, reaches peak intensity around P8, and declines to adult levels by P16. The areal and laminar patterns of the transient band of AChE activity in temporal cortex correspond to the patterns of anterograde transneuronal labeling of geniculocortical terminals following injection of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP) into the inferior colliculus. Placement of lesions that include the medial geniculate nucleus or the geniculocortical axons results in a marked decrease in AChE staining in thalamorecipient layers of auditory cortex. Placement of lesions that include the medial globus pallidus reduce AChE staining of some axons in temporal cortex of developing rats, but the dense band of AChE in layers III and IV remains. Placement of lesions in the inferior colliculus in newborn animals results in marked decrease in AChE staining in cells of the ipsilateral ventral medial geniculate nucleus and in ipsilateral auditory cortex of developing pups. These data indicate that transiently expressed AChE activity is characteristic of geniculocortical neurons, including their somata in the medial geniculate body and their terminal axons in primary auditory cortex. This AChE activity is expressed early in postnatal development, probably during the time when thalamocortical axons are proliferating in cortical layer IV and forming synaptic contacts with cortical neurons.

Acetylcholinesterase

Sensitization induced receptive field plasticity in the auditory cortex is independent of CS-modality.

Sensitization training with an auditory stimulus produces a general increase in response magnitude across the entire receptive field (RF) of neurons in the primary auditory cortex of the guinea pig (Bakin, J.S. and Weinberger, N.M., Brain Res., 536 (1990) 271-286). To determine if this effect reflects an auditory system-specific process or is caused by a process independent of the training stimulus modality, RFs in primary auditory cortex were characterized before and immediately after adult guinea pigs were given sensitization training with either an auditory or a visual training stimulus. General increases in auditory response magnitude across the RF were observed in 7 out of 7 auditory sensitization cases and 4 out of 5 visual sensitization cases. There were no statistical differences between the effects of auditory and visual sensitization training. These findings indicate that the general increases observed following sensitization training are the result of processes independent of CS modality, in contrast to the highly specific RF modifications that are caused by classical conditioning. The findings suggest that the 2 forms of RF plasticity, CS-specific re-tuning due to associative conditioning and polymodal general increases in gain due to non-associative sensitization, may reflect neural mechanisms involved in selective attention and vigilance, respectively.

Acoustic Stimulation

Primary auditory cortex in the ferret (Mustela putorius): neural response properties and topographic organization.

Microelectrode recording studies were made of the crown of the ectosylvian auditory cortex of barbiturate-anesthetized ferrets, using calibrated, sealed acoustic stimulus delivery systems. We confirmed our previous finding using free-field stimuli that this region of the ferret's cerebral cortex contains a tonotopically organized field in which neurons are briskly excited by the onset of tonal stimuli. The vast majority of neurons in this field were narrowly tuned to tone pulse frequency, with Q factors comparable to those described for cortical cells in other species. The distribution of minimum tone thresholds across the frequency representation of this field paralleled the behavioral audiogram in the same species. The majority of neurons received input from both ears, and the natures of the influences exerted by tonal stimuli at the two ears were similar in form to those described for other carnivores. The various binaural cell types had characteristic frequencies that spanned the entire tonal spectrum represented in the cortex. Preliminary mapping studies revealed that there may be a topographic segregation of cells according to their binaural interactions. The basic properties of cells in this region of the ferret's cerebrum are similar to those previously described for the cat's primary auditory cortex. The data on the binaural properties of these cells, and the spatial distribution of those cells, provide the first evidence in a species other than the cat for segregated binaural representation in the auditory cortex.

Acoustic Stimulation

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

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

The auditory cortex of the mouse: connections of the ultrasonic field.

The cortical and subcortical connections of the ultrasonic field (UF) of the auditory cortex of the house mouse (Mus musculus) were studied by using retrograde and anterograde transport of horseradish peroxidase (HRP). Small amounts of HRP were locally injected into the electrophysiologically defined UF. Superficial (layer I-IV) and deep (layer IV-VI) injections were prepared. Superficial injections led to labelling of both cells (retrograde) and terminals (anterograde) in areas of the ipsilateral primary and secondary auditory cortex and in its dorsoposterior field, in an ipsilateral dorsal association area (patches of label), probably in ipsilateral secondary somatosensory cortex, in the contralateral homotopic UF, and in the ipsilateral medial geniculate body (MGBv, MGBd, and MGBm) and caudal posterior nucleus complex. Deep injections showed the same connectivities as superficial ones and, in addition, terminals in the very caudal caudatoputamen, in the nucleus limitans and the nucleus reticularis of the thalamus, in the rostral pole, the dorsomedial, and lateral nucleus of the inferior colliculus, in the stratum griseum intermediale of the superior colliculus, and in a pontine nucleus ventromedial of the lateral lemniscus. All these projections occurred only ipsilaterally. The majority of connections, except those with the nucleus limitans, superior colliculus and pontine nucleus, suggest that UF is part of the primary anditory cortex (AI) and/or of the anterior anditory field (AAF) of the auditory cortex. Since UF has no regular tonotopy, this has important implications for the functional role that AI/AAF can have in communication-sound analysis.

Animals

Basal forebrain stimulation modifies auditory cortex responsiveness by an action at muscarinic receptors.

We have hypothesized that auditory cortex plasticity involves modification of thalamocortical transmission by basal forebrain (BF) cholinergic neurons, and that this action may involve muscarinic receptors. In a first test of this hypothesis, we report that BF stimulation can suppress or facilitate, depending on the intensity of stimulation, auditory cortical responses elicited by thalamic stimulation. BF-mediated facilitation is antagonized by atropine, implicating muscarinic receptors. These data suggest that BF cholinergic neurons functionally modify auditory cortex by regulating thalamocortical transmission.

Animals

Effects of unilateral ablation of auditory cortex on monaural cat's ability to localize sound.

1. Cats with one cochlea destroyed were trained to localize sound. After behavioral measures of the animal's accuracy of localization were made, cortical auditory areas were ablated unilaterally. 2. The results showed: a) like binaural localization, monaural localization of sound in space, as measured by the ability of an animal to move toward a sound source, depends on integrity of auditory cortex; b) it is only ablation of cortex contralateral to the functional ear that seriously affects localizing behavior; ablation of cortex ipsilateral to the intact cochlea has little or no effect on localizing behavior. 3. To explain the results, we suggest that auditory cortex is essential for an organized perception of space including the relation of the animal's position to other objects in space. We also suggest that auditory cortex contralateral to a given ear is necessary in order for the animal to recognize that a stimulus is presented to that ear of, when both ears are intact, to recognize that the stimulus to the given ear differs in some way (intensity, time of arrival, sequential arrangement of sounds) from the stimulus to the opposite ear.

Animals

The laminar organization of the prospective auditory cortex in the human fetus (11--13.5 weeks of gestation).

The prospective auditory cortex was analysed in human fetuses at 11--13,5 post-ovulatory weeks with Nissl, Golgi and E.M. techniques. At 11--12 weeks, marginal, cortical plate, intermediate, subventricular and ventricular layers were recognized. Post-migratory neurons with developing dendrites were seen in marginal layer, cortical plate and superficial part of the intermediate zone only. At 12--13,5 weeks the superficial part of the intermediate zone is transformed into the true cortical layer--"subplate layer"--characterized by maturing neurons with growing dendrites, fine axonal arborization and low cell density. The neuronal circuitry elements are thus present very early in the prospective auditory cortex and distributed throughout the deep cortical plate of the "subplate layer" corresponding to the synaptic territory of other areas of the human fetal cortex.

Auditory Cortex

Magnetic responses of the human auditory cortex to noise/square wave transitions.

We recorded evoked magnetic fields from the human auditory cortex to noise/square wave sequences. Two prominent deflections were observed: one 100 msec after the noise onset (N100m) and another 100 msec after the noise/square wave transition (N100m'). The amplitude of N100m' increased with decrease in square wave frequency from 2 kHz to 0.125 kHz and with increase in square wave duration from 4 msec to 200 msec. The latency of N100m' was on the average 23 msec longer for noise durations of 60 msec than 310 msec, whereas the amplitude of N100m' did not change. Increase in interstimulus interval from 1.1 to 8.8 sec enhanced the amplitude of N100m significantly more than that of N100m'. The small interaction between N100m and N100m' and their different recovery cycles suggest that different activation patterns underlie these two 100 msec responses at the auditory cortex.

Adult

[A histochemical study of acetylcholinesterase in intact and deafferented cat auditory cortex].

The peculiarities of the AChE distribution were investigated in the intact cat auditory cortex and during early period of its neuronal isolation. It is shown that in the isolated cortex slab the staining of the AChE containing fibre disappeared from the neuropile, while in the intact cortex it was well pronounced. AChE accumulation was observed in the proximal parts of the transsected thalamo-cortical fibres. It is supposed that the AChE-containing fibres in the auditory cortex belong to nonspecific thalamic inputs.

Acetylcholinesterase

Reactions of human auditory cortex to a change in tone duration.

Auditory evoked magnetic fields were measured using a stimulation sequence where repetitive identical tone bursts were randomly and infrequently replaced by shorter tones. The deviation in the stimulus duration evoked a specific response, the mismatch field, peaking about 130 ms after the end of the shorter stimulus. The response can be explained by neural activity at the supratemporal auditory cortex. The results suggest the existence of a neural mechanism specific to changes in sound duration.

Acoustic Stimulation

[Comparative study of interneuronal relations in the auditory cortex of awake and anesthetized cats].

The character of interneuronal relations in the auditory cortex of alert and anaesthetized cats (nembutal) with chronicly inplanted electrodes was studied with the method of statistic analysis of cross-intervals of the two impulse series. The analysis of the histograms, obtained by means of processing a neuronal activity, showed that nembutal did not eliminate the dependent relations between neurones and that in the majority of cases the types of these relations are either retained or supplemented with new components. Experiments with a reduced dose of nembutal permitted to trace in time the changes in the amount of the inhibitory and excitatory interrelations in the anaesthetized state, and to compare these changes to the changes in the frequency of spike activity. It was found that nembutal predominantly suppresses the activity of the neurones, generating small spikes. The number of inhibitory connections is reduced simultaneously. Such synchroneity permits to assume the participation of the neurones generating small spikes in the establishment of inhibitory interrelations in the cat auditory cortex.

Anesthesia, General

Role of context in the expression of learning-induced plasticity of single neurons in auditory cortex.

Classical conditioning produces frequency-specific plasticity of receptive fields (RFs) of single neurons in cat auditory cortex (Diamond & Weinberger, 1986). In this article we show that although plasticity may be observed during both training trials and determination of RFs, it is usually expressed in a qualitatively different form (e.g., decreased response during conditioning vs. increased response to this same conditioned stimulus in the postconditioning RF). This differential expression of learning-induced plasticity provides evidence for a role of context in neurophysiological mechanisms of learning in auditory cortex. A model of cortical neurons functioning within a mosaic of influences is presented. The Functional Mosaic model views the induction and expression of plasticity as separate processes.

Animals

Extrathalamic ascending projections to physiologically identified fields of the cat auditory cortex.

The neurons of origin of ascending extrathalamic projections to the auditory cortex were labeled retrogradely with WGA-HRP injected in physiologically identified auditory cortical fields of the cat (anterior (AAF), primary (AI), posterior (PAF) and secondary (AII) fields). After injection in the tonotopically organized auditory cortical fields (AAF, AI and PAF), labeled neurons were distributed in 7 extrathalamic subcortical regions included in one or the other of 2 distinct systems of ascending projections to the neocortex. In the 'diffuse' system of projection, labeled neurons were observed bilaterally in the locus coeruleus, the nuclei of the raphe, the lateral hypothalamus, ipsilaterally in the ventromedial mesencephalic tegmentum and the basal forebrain; in the 'accessory sensory' system of projection, labeled neurons were found ipsilaterally in the nucleus of the brachium of the inferior colliculus and bilaterally in the claustrum. After injection in AII, labeled neurons were seen only in the 'diffuse' system of projection. For AAF and AI, the major contribution to the total extrathalamic ascending input originated from the lateral hypothalamus, whereas for AII it was the locus coeruleus. In contrast, PAF received extrathalamic ascending inputs mainly from the claustrum. Anterogradely labeled corticofugal terminal fields were found only in the nucleus of the brachium of the inferior colliculus and, after injection in PAF, in the claustrum.

Acetylcholine