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PubMed2001

Human primary auditory cortex: cytoarchitectonic subdivisions and mapping into a spatial reference system.

The transverse temporal gyrus of Heschl contains the human auditory cortex. Several schematic maps of the cytoarchitectonic correlate of this functional entity are available, but they present partly conflicting data (number and position of borders of the primary auditory areas) and they do not enable reliable comparisons with functional imaging data in a common spatial reference system. In order to provide a 3-D data set of the precise position and extent of the human primary auditory cortex, its putative subdivisions, and its topographical intersubject variability, we performed a quantitative cytoarchitectonic analysis of 10 brains using a recently established technique for observer-independent definition of areal borders. Three areas, Te1.1, Te1.0, and Te1.2, with a well-developed layer IV, which represent the primary auditory cortex (Brodmann area 41), can be identified along the mediolateral axis of the Heschl gyrus. The cell density was significantly higher in Te1.1 compared to Te1.2 in the left but not in the right hemisphere. The cytoarchitectonically defined areal borders of the primary auditory cortex do not consistently match macroanatomic landmarks like gyral and sulcal borders. The three primary auditory areas of each postmortem brain were mapped to a spatial reference system which is based on a brain registered by in vivo magnetic resonance imaging. The integration of a sample of postmortem brains in a spatial reference system allows one to estimate the spatial variability of each cytoarchitectonically defined region with respect to this reference system. In future, the transfer of in vivo structural and functional data into the same spatial reference system will enable accurate comparisons of cytoarchitectonic maps of the primary auditory cortex with activation centers as established with functional imaging procedures.

Adult↗
PubMed2003

Single electrode micro-stimulation of rat auditory cortex: an evaluation of behavioral performance.

A combination of electrophysiological mapping, behavioral analysis and cortical micro-stimulation was used to explore the interrelation between the auditory cortex and behavior in the adult rat. Auditory discriminations were evaluated in eight rats trained to discriminate the presence or absence of a 75 dB pure tone stimulus. A probe trial technique was used to obtain intensity generalization gradients that described response probabilities to mid-level tones between 0 and 75 dB. The same rats were then chronically implanted in the auditory cortex with a 16 or 32 channel tungsten microwire electrode array. Implanted animals were then trained to discriminate the presence of single electrode micro-stimulation of magnitude 90 microA (22.5 nC/phase). Intensity generalization gradients were created to obtain the response probabilities to mid-level current magnitudes ranging from 0 to 90 microA on 36 different electrodes in six of the eight rats. The 50% point (the current level resulting in 50% detections) varied from 16.7 to 69.2 microA, with an overall mean of 42.4 (+/-8.1) microA across all single electrodes. Cortical micro-stimulation induced sensory-evoked behavior with similar characteristics as normal auditory stimuli. The results highlight the importance of the auditory cortex in a discrimination task and suggest that micro-stimulation of the auditory cortex might be an effective means for a graded information transfer of auditory information directly to the brain as part of a cortical auditory prosthesis.

Animals↗
PubMed2003

Spectrotemporal structure of receptive fields in areas AI and AAF of mouse auditory cortex.

The mouse is a promising model system for auditory cortex research because of the powerful genetic tools available for manipulating its neural circuitry. Previous studies have identified two tonotopic auditory areas in the mouse-primary auditory cortex (AI) and anterior auditory field (AAF)- but auditory receptive fields in these areas have not yet been described. To establish a foundation for investigating auditory cortical circuitry and plasticity in the mouse, we characterized receptive-field structure in AI and AAF of anesthetized mice using spectrally complex and temporally dynamic stimuli as well as simple tonal stimuli. Spectrotemporal receptive fields (STRFs) were derived from extracellularly recorded responses to complex stimuli, and frequency-intensity tuning curves were constructed from responses to simple tonal stimuli. Both analyses revealed temporal differences between AI and AAF responses: peak latencies and receptive-field durations for STRFs and first-spike latencies for responses to tone bursts were significantly longer in AI than in AAF. Spectral properties of AI and AAF receptive fields were more similar, although STRF bandwidths were slightly broader in AI than in AAF. Finally, in both AI and AAF, a substantial minority of STRFs were spectrotemporally inseparable. The spectrotemporal interaction typically appeared in the form of clearly disjoint excitatory and inhibitory subfields or an obvious spectrotemporal slant in the STRF. These data provide the first detailed description of auditory receptive fields in the mouse and suggest that although neurons in areas AI and AAF share many response characteristics, area AAF may be specialized for faster temporal processing.

Acoustic Stimulation↗
PubMed2000

Frequency-dependent responses exhibited by multiple regions in human auditory cortex.

Recordings in experimental animals have detailed the tonotopic organization of auditory cortex, including the presence of multiple tonotopic maps. In contrast, relatively little is known about tonotopy within human auditory cortex, for which even the number and location of tonotopic maps remains unclear. The present study begins to develop a more complete picture of cortical tonotopic organization in humans using functional magnetic resonance imaging, a technique that enables the non-invasive localization of neural activity in the brain. Subjects were imaged while listening to lower- (below 660 Hz) and higher- (above 2490 Hz) frequency stimuli presented alternately and at moderate intensity. Multiple regions on the superior temporal lobe exhibited responses that depended upon stimulus spectral content. Eight of these 'frequency-dependent response regions' (FDRRs) were identified repeatedly across subjects. Four of the FDRRs exhibited a greater response to higher frequencies, and four exhibited a greater response to lower frequencies. Based upon the location of the eight FDRRs, a correspondence is proposed between FDRRs and anatomically defined cortical areas on the human superior temporal lobe. Our findings suggest that a larger number of tonotopically organized areas exist (i.e., four or more) in the human auditory cortex than was previously recognized.

Acoustic Stimulation↗
PubMed1978

Response variability in the mammalian auditory cortex: an objection to feature detection?

Research strategy in the auditory system has tended to parallel that in the visual system, where neurons have been shown to respond selectively to specific stimulus parameters. Auditory neurons have been shown to be sensitive to changes in acoustic parameters, but only rarely have neurons been reported that respond exclusively to only one biologically significant sound. Even at higher levels of the auditory system very few cells have been found that could be described as "vocalization detectors." In addition, variability in responses to artificial sounds have been reported for auditory cortical neurons similar to the response variability that has been reported in the visual system. Recent evidence indicates that the responses of auditory cortical neurons to species-specific vocalizations can also be labile, varying in both strength and selectivity. This is especially true of the secondary auditory cortex. This variability, coupled with the lack of extreme specificity in the secondary auditory cortex, suggests that secondary cortical neurons are not well suited for the role of "vocalization detectors."

Acoustic Stimulation↗
PubMed2001

Spectral and temporal processing in human auditory cortex.

We used positron emission tomography to examine the response of human auditory cortex to spectral and temporal variation. Volunteers listened to sequences derived from a standard stimulus, consisting of two pure tones separated by one octave alternating with a random duty cycle. In one series of five scans, spectral information (tone spacing) remained constant while speed of alternation was doubled at each level. In another five scans, speed was kept constant while the number of tones sampled within the octave was doubled at each level, resulting in increasingly fine frequency differences. Results indicated that (i) the core auditory cortex in both hemispheres responded to temporal variation, while the anterior superior temporal areas bilaterally responded to the spectral variation; and (ii) responses to the temporal features were weighted towards the left, while responses to the spectral features were weighted towards the right. These findings confirm the specialization of the left-hemisphere auditory cortex for rapid temporal processing, and indicate that core areas are especially involved in these processes. The results also indicate a complementary hemispheric specialization in right-hemisphere belt cortical areas for spectral processing. The data provide a unifying framework to explain hemispheric asymmetries in processing speech and tonal patterns. We propose that differences exist in the temporal and spectral resolution of corresponding fields in the two hemispheres, and that they may be related to anatomical hemispheric asymmetries in myelination and spacing of cortical columns.

Acoustic Stimulation↗
PubMed1987

Different analysis of frequency and amplitude modulations of a continuous tone in the human auditory cortex: a neuromagnetic study.

We have measured auditory evoked magnetic fields to intermittent frequency and amplitude modulations (FMs and AMs) of a continuous tone in 6 healthy humans. The stimuli were presented in pairs separated by 500 ms in four different combinations (FM-AM, FM-FM, AM-FM and AM-AM). Both modulations elicited neuromagnetic responses of similar waveforms: the largest deflection, N100m (magnetic counterpart of the electric N100), was preceded by a low amplitude P60m and followed by P200m. For stimuli of different types, the decrease of N100m from the first to the second response was less than expected from the recovery cycle of the responses, estimated from the pairs of similar stimuli. We interpret these results as evidence for different processing of amplitude and frequency modulations in the auditory pathways up to the level of supratemporal auditory cortex.

Auditory Cortex↗
PubMed1989

Functional subdivisions in the auditory cortex of the guinea pig.

The auditory fields in the cortex of the guinea pig were investigated with microelectrode mapping techniques. Pure tones of varying frequencies and amplitudes were used as acoustic stimuli. Mainly, multiunit activity was recorded. A large tonotopic area is found in the anterior half of the auditory cortex. This area is named the anterior field (field A). Frequency tuning curves of multiunits in field A are generally narrow. Responses to tone stimuli are strong, and latencies are short. Low best frequencies are represented rostrally, high best frequencies caudally. The tonotopy is continuous and quite regular. Field A is narrow dorsally and becomes gradually broader ventrally. Correspondingly, the isofrequency lines slightly diverge from dorsal to ventral. Caudal to the first field, there is a second, smaller tonotopic area. It lies in the dorsal half of the posterior auditory cortex and is therefore named the dorsocaudal field (field DC). The frequency specificity of the cell clusters in this area is as strong as in field A, but the tonotopy is discontinuous: In the dorsal half of field DC, high best frequencies (16-32 kHz) are represented rostrally; the low frequencies (0.5-2.8 kHz) are represented immediately caudal to the high frequencies, while the intermediate frequencies are missing. Ventrally in field DC, the frequency representation is more complete. Except for this discontinuous map, we did not notice any differences between fields A and DC. A third tonotopic field was found rostral to field A. This field extends over a surface of less than 1 mm2 and was named the small field (field S). It contains a complete representation of the frequency range; high best frequencies are located rostrally, low frequencies caudally. The response latencies are slightly longer in field S than in fields A or DC, and the tuning curves are broader. A broad strip of nontonotopic cortex (auditory belt) surrounds fields A and DC caudally. We subdivided this area into the dorsocaudal and the ventrocaudal belt region. In both areas, tuning curves are often broad, and response latencies are longer than in the tonotopic cortex. In the dorsocaudal belt, most multiunits react with a phasic on-response to pure tones; in the ventrocaudal belt, tonic responses occur more frequently. Another nontonotopic region is located in the anterior auditory cortex, rostral to the tonotopic fields, and was therefore named the rostral belt. Tuning curves in this area are broad, latencies are short, and response thresholds are often high. In the discussion, the guinea pig is compared with other mammalian species.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗
PubMed2001

Relating cluster and population responses to natural sounds and tonal stimuli in cat primary auditory cortex.

Most information about neuronal properties in primary auditory cortex (AI) has been gathered using simple artificial sounds such as pure tones and broad-band noise. These sounds are very different from the natural sounds that are processed by the auditory system in real world situations. In an attempt to bridge this gap, simple tonal stimuli and a standard set of six natural sounds were used to create models relating the responses of neuronal clusters in AI of barbiturate-anesthetized cats to the two classes of stimuli. A significant correlation was often found between the response to the separate frequency components of the natural sounds and the response to the natural sound itself. At the population level, this correlation resulted in a rate profile that represented robustly the spectral profiles of the natural sounds. There was however a significant scatter in the responses to the natural sound around the predictions based on the responses to tonal stimuli. Going the other way, in order to understand better the non-linearities in the responses to natural sounds, responses of neuronal clusters were characterized using second order Volterra kernel analysis of their responses to natural sounds. This characterization predicted reasonably well the amplitude of the response to other natural sounds, but could not reproduce the responses to tonal stimuli. Thus, second order non-linear characterizations, at least those using the Volterra kernel model, do not interpolate well between responses to tones and to natural sounds in auditory cortex.

Acoustic Stimulation↗
PubMed1983

Connections of the auditory cortex of the brush-tailed possum, Trichosurus vulpecula.

The afferent connections of the auditory cortex of the acallosal marsupial Trichosurus vulpecula have been studied using the retrograde transport of horseradish peroxidase. The enzyme was applied by electrophoresis from a micropipette previously used to define physiologically the organization of the auditory cortex. Three major (lateral, dorsomedial and ventromedial) sectors of the medial geniculate contribute axons to the ipsilateral auditory cortex; the connections from at least the lateral sector are topographically arranged. Axons reach the auditory field from the contralateral cortex by way of the anterior commissure and fasciculus aberrans.

Afferent Pathways↗
PubMed1998

Salicylate and quinine selectively increase spontaneous firing rates in secondary auditory cortex.

This study presents firing rates for simultaneously recorded spontaneous and stimulus driven multi-unit activity in primary auditory cortex (AI), anterior auditory field (AAF) and secondary auditory cortex (AII) in cats before and after application of salicylate or quinine. From 21 cats, in three cortical areas simultaneously, a total of 1533 multi-unit files were obtained. The data suggest (1) that both salicylate and quinine significantly increase spontaneous firing rates in AII, whereas in AI and AAF both quinine and salicylate reduced the spontaneous rate; (2) the effect of both drugs was to increase spontaneous rates for recording sites with high characteristic frequency (CF) and a tendency to decrease them for low CF sites; (3) the mean stimulus driven firing rates were not affected by either drug except for a decrease produced by quinine in AI; (4) changes in driven firing rate were positively correlated with changes in spontaneous firing rates. This suggests that tinnitus inducing agents selectively increase spontaneous firing rates in the extralemniscal pathway.

Acoustic Stimulation↗
PubMed1983

The inability of squirrel monkeys to localize sound after unilateral ablation of auditory cortex.

The ability of squirrel monkeys to localize brief sounds was tested before and after unilateral ablation of auditory cortex. Results indicated that, after unilateral ablation of auditory cortex, monkeys could no longer localize sound accurately in the hemifield opposite to the side of lesion. These data demonstrate that contralaterally-directed functioning of auditory cortex occurs in primates.

Animals↗
PubMed1997

Fear conditioning enhances different temporal components of tone-evoked spike trains in auditory cortex and lateral amygdala.

Single neurons were recorded in freely behaving rats during fear conditioning from areas of auditory cortex that project to the lateral nucleus of the amygdala (LA). The latency and rate of conditioning and extinction were analyzed, and the results were compared to previous recordings from LA itself. Auditory cortex neurons took more trials to learn, and they responded more slowly than LA neurons within trials. Short-latency plasticity in LA, therefore, reflects inputs from the auditory thalamus rather than the auditory cortex. Unlike LA cells, some auditory cortex cells showed late conditioned responses that seemed to anticipate the unconditioned stimulus, while others showed extinction-resistant memory storage. Thus, rapid conditioning of fear responses to potentially dangerous stimuli depends on plasticity in the amygdala, while cortical areas may be particularly involved in higher cognitive (mnemonic and attentional) processing of fear experiences.

Acoustic Stimulation↗
PubMed1996

Recovery of function after neonatal ablation of the auditory cortex in rats (Rattus norvegicus).

Functional recovery following neonatal ablation of the auditory cortex was surveyed in 28 rats. Fourteen neonatal-lesioned rats had their temporal cortex lesioned on the date of birth (P1); 7 adult-lesioned rats had their auditory cortex lesioned at P60; and 7 rats served as controls. The training consisted of two phases using task with a Go/No Go procedure. The first task was to detect the presentation of a tone pulse. The second task was to discriminate the pulse rate of a tone pulse. Results showed that the achievements of neonatal-lesioned and control groups were comparable in both tasks. Adult-lesioned rats, however, failed to discriminate temporal patterns, although they could detect tone presentation as efficiently as the other two groups. These findings suggested that discrimination of temporal patterns could be a critical function of the auditory cortex and that brain injury in infancy was more compensated than the comparable damage in adulthood. Neurological plasticity was suggested in the recovery of function in our neonatal-lesioned animals.

Acoustic Stimulation↗
PubMed1978

Auditory cortex lesions and discrimination of spatial location by the rat.

Five normal rats and four rats with bilateral lesions of auditory cortex were tested by the conditioned suppression procedure to determine their abilities to discriminate between spatially separated sound sources. The discrimination involved detection of a change in location of a train of clicks from a speaker on the animals' left to a speaker on the right. The separation between speakers was varied from 180 degrees to 90 degrees, 45 degrees, 22 degrees, 12 degrees, 6 degrees, and psychophysical functions were obtained using a method of descending limits. Both normal and brain-damaged animals were capable of discriminating left from right clicks and psychophysical curves were similar for the two groups. Histological analysis indicated that the lesions in each of the four brain-damaged rats destroyed primary auditory cortex as well as surrounding belt areas. Therefore, for the rat, auditory cortex was not found to be essential for discrimination of the spatial locations of auditory stimuli. The results are discussed in light of impairments in sound localization following lesions of auditory cortex in other mammalian species.

Acoustic Stimulation↗
PubMed1986

Corticocortical connections of cat primary auditory cortex (AI): laminar organization and identification of supragranular neurons projecting to area AII.

The laminar distribution and structure of the supragranular cells projecting from primary auditory cortex (AI) to the second auditory cortex (AII) in the cat were studied with horseradish peroxidase. Injections in AII retrogradely labeled somata in ipsilateral cortical layers I-VI of AI. A bimodal laminar disposition was found, with approximately 40% of the labeled cells in layer III, 25% in layer V, and 10-15% each in layers II, IV, and VI; only a few cells were found in layer I. The labeled cells were scattered in small aggregates between which unlabeled neurons were interspersed. There was some, though not a strict, topographical distribution of the labeled cells according to the locus of the injection in AII. Injections in the caudal part of AII labeled cells in more rostral AI, while rostral AII injections labeled cells in more caudal AI. Ventral AII injections labeled more ventrally located AI cells, while more dorsal AII injections labeled more dorsally situated AI cells. AII injections also labeled cells in other auditory cortex subdivisions, including the posterior ectosylvian, ventroposterior, temporal, and dorsal auditory zone/suprasylvian fringe cortical areas, and in some non-auditory cortical areas. In layers II and III, both pyramidal and non-pyramidal cells were labeled. More pyramidal cells were labeled in layer III than layer II (80% vs. 62%), and the proportion of non-pyramidal cells in layer II was more than twice that in layer IV (27% vs. 12%). The types of labeled cells were distinguished from one another on the basis of size, somatic and dendritic shape, and laminar distribution. The profiles of labeled cells in these experiments were compared to, and correlated with, those in Golgi-impregnated material. In layer II, the classes of corticocortical projecting cells consisted of small and medium-sized pyramidal, bipolar, and multipolar cells. Those in layer III included small, medium-sized, and large pyramidal neurons, and bipolar and multipolar cells. The average somatic area of the labeled cells did not differ significantly from that of the unlabeled cells, and both were about equal in somatic size to neurons accumulating tritiated gamma-aminobutyric acid in layers II and III. These findings suggest that there is convergent, ipsilateral input onto AII from every layer in AI, and from other cortical auditory and non-auditory areas. A morphologically heterogeneous population of cells in AI contributes to these projections. Diversity in the cytological origins of corticocortical projections implies functional differences between layers II and III since the latter also projects commissural, while layer II in the cat, does not.

Animals↗
PubMed1996

Influence of urethane anesthesia on neural processing in the auditory cortex analogue of a songbird.

Functional maps of auditory response areas were derived from multi-unit recordings in the caudal telencephalon of the starling (Sturnus vulgaris L.). A regular grid of recording sites with distances of 200 microns horizontally and 100 microns vertically was placed over the auditory cortex analogue. Within one plane, mapping of auditory responses was first performed in the awake bird and then repeated under urethane anesthesia. The data from both experimental approaches differ considerably. Urethane reduces the spontaneous discharge rate significantly. Under anesthesia, inhibition decreases in all auditory subunits. Excitation is less affected. Eight auditory subcenters were divided into three groups according to the changes in their excitatory responses. In the first group 'on' and sustained excitation changed only weakly. These areas are thought to receive direct inputs from the diencephalon. In the second group, 'on' and sustained excitation are substantially reduced. These subcenters seem to receive projections from other forebrain areas. In subunits of the third group, an increase in sustained excitation is correlated to a decrease in inhibition. Within some specific centers, distinct natural calls, for example the bird's own song, elicit stronger responses under anesthesia than other stimuli.

Acoustic Stimulation↗
PubMed1980

[Quantitative and qualitative characteristics of synapses in different layers of the auditory cortex].

The electron-microscopic examination of synapses in different layers of the cat auditory cortex was performed. 53% of them were located on dendritic spines, 37% on dendrites and 10% on neuronal soma. All synapses were separated into type I and type II according to Gray. The type I synapses amounted to 91% (69.5% were of type Ia and 21.5% of type Ib), the type II synapses constituted 9%. The type I synapses were located mainly on dendrites and dendritic spines, the type II--on neuronal soma, axon hillocks and large dendrites. 60 h after complete neuronal isolation of a portion of the auditory cortex 22.8% of synapses revealed signs of degeneration. No type II degenerating synapses were found. This indicates that they are formed by axons of intracortical neurons. Quantitative and qualitative composition of synapses in different layers of the auditory cortex is shown to be different.

Animals↗