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PubMed2000

Mechanisms and streams for processing of "what" and "where" in auditory cortex.

The functional specialization and hierarchical organization of multiple areas in rhesus monkey auditory cortex were examined with various types of complex sounds. Neurons in the lateral belt areas of the superior temporal gyrus were tuned to the best center frequency and bandwidth of band-passed noise bursts. They were also selective for the rate and direction of linear frequency modulated sweeps. Many neurons showed a preference for a limited number of species-specific vocalizations ("monkey calls"). These response selectivities can be explained by nonlinear spectral and temporal integration mechanisms. In a separate series of experiments, monkey calls were presented at different spatial locations, and the tuning of lateral belt neurons to monkey calls and spatial location was determined. Of the three belt areas the anterolateral area shows the highest degree of specificity for monkey calls, whereas neurons in the caudolateral area display the greatest spatial selectivity. We conclude that the cortical auditory system of primates is divided into at least two processing streams, a spatial stream that originates in the caudal part of the superior temporal gyrus and projects to the parietal cortex, and a pattern or object stream originating in the more anterior portions of the lateral belt. A similar division of labor can be seen in human auditory cortex by using functional neuroimaging.

Acoustic Stimulation↗
PubMed2005

Interfield differences in intensity and frequency representation of evoked potentials in rat auditory cortex.

Existing studies have demonstrated interfield differences in functional organizations and neuronal responsive properties at a single neuron level in the auditory cortex, suggesting complicated encoding of sound frequency and intensity. The objective of present work is, by characterizing cortical auditory evoked potentials (AEPs), to bridge neural characteristics between a single neuron and field levels and to identify the interfield differences in the auditory cortex specifically in terms of spatial representation, which will be useful in guiding future unit studies. The AEP mapping found that each of auditory fields, which could be identified by a different tonotopic representation, showed interfield differences in an intensity-dependent spatial change, amplitude, latency, and amplitude-SPL (sound pressure level) function. These results also showed that many aspects of cortical representation were based on the cochlear properties, yet some were inconsistent. The intensity-dependent shift of activation in AI paralleled the tonotopic axis, which was similar to the place code in cochlea, while the shift in AAF and VAF did not parallel. Nevertheless, the amplitude-SPL function suggested that an underlying mechanism of all these shifts can be a compressive nonlinearity to CF tone, which is possibly formed in the cochlea and still preserved in the cortex. These results suggest that each field modifies the representation to handle a different aspect of sound information, which can be better analyzed than the cochlear representation.

Acoustic Stimulation↗
PubMed2006

Binaural-level functions in ferret auditory cortex: evidence for a continuous distribution of response properties.

Many previous studies have subdivided auditory neurons into a number of physiological classes according to various criteria applied to their binaural response properties. However, it is often unclear whether such classifications represent discrete classes of neurons or whether they merely reflect a potentially convenient but ultimately arbitrary partitioning of a continuous underlying distribution of response properties. In this study we recorded the binaural response properties of 310 units in the auditory cortex of anesthetized ferrets, using an extensive range of interaural level differences (ILDs) and average binaural levels (ABLs). Most recordings were from primary auditory fields on the middle ectosylvian gyrus and from neurons with characteristic frequencies >5 kHz. We used simple multivariate statistics to quantify a fundamental coding feature: the shapes of the binaural response functions. The shapes of all 310 binaural response surfaces were represented as points in a five-dimensional principal component space. This space captured the underlying shape of all the binaural response surfaces. The distribution of binaural level functions was not homogeneous because some shapes were more common than others. Despite this, clustering validation techniques revealed no evidence for the existence of discrete, or partially overlapping, clusters that could serve as a basis for an objective classification of binaural-level functions. We also examined the gradients of the response functions for the population of units; these gradients were greatest near the midline, which is consistent with free-field data showing that cortical neurons are most sensitive to changes in stimulus location in this region of space.

Acoustic Stimulation↗
PubMed1993

Basal forebrain stimulation facilitates tone-evoked responses in the auditory cortex of awake rat.

The effects of unilateral basal forebrain stimulation on the tone-evoked responses recorded in the auditory cortex ipsilateral and contralateral to the stimulation site, were investigated in fully awake rats. After 10 tone alone presentations, 20 pairing trials were given during which the basal forebrain stimulation was followed by the tone 30 ms later. Ten test-tones were presented immediately, 15 min and 1 h after pairing. Immediately after pairing, the short-latency "on" and "off" tone-evoked responses were enhanced in the ipsilateral but not in the contralateral cortex. This enhancement did not persist 15 min later. Systemic atropine injection prevented the ipsilateral facilitation. The responses to the tone were not modified when tested after 20 basal forebrain stimulations delivered in the absence of the tone. These results are the first demonstration in awake animals that an activation of the auditory cortex by cholinergic neurons of the basal forebrain is able to facilitate cortical responsiveness. A temporal contiguity between the cholinergic activation and the neuronal discharges elicited by the sensory stimulus is required for the facilitation to take place. The results are compared to previous ones obtained in anesthetized animals, and the functional role of cholinergic activation from the basal forebrain in cortical processing is discussed.

Acoustic Stimulation↗
PubMed1984

[Proposed inhibitory neurons of the auditory cortex in the cat].

A complex of properties of inhibitory neurons in the auditory cortex is defined according to the peculiarities of IPSPs recorded in many cortical neurons. 12 units with such properties were found in a group consisting of 54 cells. These low-threshold units were localized predominantly in layers III-IV, had no background activity and responded to afferent volleys by short-latent barbiturate-resistant discharges (1-4 spikes).

Animals↗
PubMed1996

On determinants of first-spike latency in auditory cortex.

The first-spike latency of neurones at any level of the auditory pathway decreases with stimulus amplitude. As stimuli are generally shaped with rise functions to avoid spectral splatter, a common interpretation of the latency decrease is that the amplitude of the signal reaches the neurone's firing threshold earlier during the rise time. We demonstrate here, for auditory cortex neurones and by varying the amplitude and rise time of tonal stimuli, that this threshold model is inadequate to account for the observed latency changes, particularly when adaptive processes are taken into account. The data raise the possibility that latency may be a function of other properties associated with a signal's onset, such as rate of change of peak pressure.

Acoustic Stimulation↗
PubMed2001

Asymmetrical localization of benzodiazepine receptors in the human auditory cortex.

In humans, administration of benzodiazepines (BZD) has been shown to have an asymmetrical effect on the medial olivocochlear system. Indeed, a decrease of evoked otoacoustic emission suppression by contralateral acoustic stimulation, which explores the medial olivocochlear efferent system, was observed in the right ear, with no left ear effect. This result suggests a possible left-right auditory pathway BZD receptor asymmetry. Given the anatomical link between auditory centers and the medial olivocochlear system, the existence of a larger volume of cortical connecting fibers in the left hemisphere, and the possible link between BZD receptor density and neuronal density, we tested the hypothesis of an asymmetrical localization of BZD receptors in the auditory system in 10 right-handed subjects using [11C]flumazenil positron emission tomography. Semi-quantitative measurements of flumazenil binding were evaluated in Heschl's gyrus showing a left-right asymmetry in favor of left auditory cortex. This result indicates a higher density of neurons in left auditory cortex. The possible link between neurochemical asymmetry and functional asymmetry, and the perceptual outcome of BZD administration, will be discussed.

Adult↗
PubMed2003

Early and late patterns of stimulus-related activity in auditory cortex of trained animals.

Epidural electrocorticograms over the right auditory cortex (field AI) were measured using implanted 18-channel (3 x 6) electrode arrays in four animals (Mongolian gerbil) trained to discriminate between a rising and a falling frequency modulated tone (frequency range 2-4 kHz). Using a previously introduced classification procedure, transient patterns of cortical activity suitable to discriminate between the rising and the falling modulation were identified. Early (locked to stimulus onset) and late (emerging at variable times poststimulus) patterns could be differentiated. Deletion of increasing numbers of randomly selected electrodes was used to determine a critical density of recording channels required to capture the discriminative power of the early and late patterns. Statistical analysis of the classification revealed a sigmoid dependence of the discriminative power from the number of remaining electrodes with an inflection point at 12 electrodes. The analysis of the minima of the classification statistic revealed that in the early patterns discriminative information was focal on regions corresponding to the tonotopic representation of the stimuli, whereas in late patterns this information seemed to be distributed nonfocally across larger cortical regions. This analysis supports the previous notion of the coexistence of topographically organized activity states related to the physical stimulus features and nontopographically organized states determined largely by intrinsic factors (Ohl et al. 2001).

Acoustic Stimulation↗
PubMed1978

Effect of auditory cortex ablation on localization and discrimination of brief sounds.

1. Dogs with bilateral auditory cortex lesions were tested on their ability to localize and discriminate brief sounds. In each test the animals were required to approach one of two goal boxes in order to indicate their response. 2. The results showed: a) that the operated animals could not solve the localization tasks when the goal boxes were located more than 125 cm away, but could solve the task if the goal boxes were located closer to the animal; b) that the operated animals could successfully discriminate brief bursts of click trains (i.e., 100/s versus 10/s, 0.3 s duration) even when required to indicate their discrimination by moving to goal boxes located 250 cm away, in spite of the fact that they could not successfully localize these sounds under similar conditions; c) that the operated animals tracked the source of a continuous sound instead of localizing it in a normal manner. 3. It appears that the deficit in sound localization resulting from cortical ablation is not due to any impairment in auditory attention or memory. Furthermore, the deficit cannot be ascribed to an inability to make a spatial response to an auditory cue. Instead, the deficit may be the result of a disconnection of the sound-localization mechanism from the motor mechanism necessary for some, though not all, behavioral responses.

Animals↗
PubMed2006

Focal projections of cat auditory cortex to the pontine nuclei.

The pontine nuclei (PN) receive projections from the auditory cortex (AC) and they are a major source of mossy fibers to the cerebellum. However, they have not been studied in detail using sensitive neuroanatomical tracers, and whether all AC areas contribute to the corticopontine (CP) system is unknown. We characterized the projection patterns of 11 AC areas with WGA-HRP. We also compared them with their corticothalamic and corticocollicular counterparts. A third objective was to analyze the structure of the CP axons and their terminals with BDA. Both tracers confirm that all AC areas projected to lateral, central, and medial ipsilateral pontine divisions. The strongest CP projections were from nontonotopic and polymodal association areas. Preterminal fibers formed single terminal fields having many boutons en passant as well as terminal endings, and there was a specific morphological pattern for each pontine target, irrespective of their areal origin. Thus, axons in the medial division had a simpler terminal architecture (type 1 terminal plexus); both the central and lateral pons received more complex endings (type 2 terminal plexus). Auditory CP topographical distribution resembled visual and somatosensory CP projections, which preserve retinotopy and somatotopy in the pons, respectively. However, the absence of pontine tonotopy suggests that the AC projection topography is unrelated to tonotopy. CP input to the medial and central pons coincides with the somatosensory and visual cortical inputs, respectively, and such overlap might subserve convergence in the cerebellum. In contrast, lateral pontine input may be exclusively auditory.

Animals↗
PubMed1992

Spatio-temporal pattern of frequency representation in the auditory cortex of guinea pigs.

The spatio-temporal pattern of sound-evoked neural activity in the guinea pig auditory cortex was studied by optical recording with the aid of voltage-sensitive dye. Changes in light intensity induced by sounds at various frequencies and pressure levels were recorded with a 12 x 12 array of photodiodes. The amplitudes of the responses were displayed as sequential two-dimensional images. Tonotopical organization was found in two subdivisions of the auditory cortex, the anterior field (field A) and the dorsocaudal field (field DC). The frequency gradients in fields A and DC had a mirror-image relationship. This agrees with results obtained by the microelectrode technique. However, the tonotopic response observed in our study was transient. The focal activity that began in field A propagated in two directions; dorsally along the iso-frequency bands in field A, and caudally toward field DC. This suggests that the sound information processing initiates at field A, and its outputs are transferred to field DC, which is probably a hierarchically higher center.

Acoustic Stimulation↗
PubMed1993

Tonotopic organization, architectonic fields, and connections of auditory cortex in macaque monkeys.

Microelectrode recordings were used to investigate the tonotopic organization of auditory cortex of macaque monkeys and guide the placement of injections of wheat germ agglutinin-horse radish peroxidase (WGA-HRP) and fluorescent dyes. Anatomical and physiological results were later related to histological distinctions in the same brains after sections were processed for cytoarchitecture, myeloarchitecture, acetylcholinesterase (AchE), or cytochrome oxidase (CO). The experiments produced several major findings. (1) Neurons throughout a broad expanse of cortex were highly responsive to pure tones, and best frequencies could be determined for neurons in arrays of recording sites. (2) The microelectrode recordings revealed two systematic representations of tone frequencies, the primary area (AI) and a primary-like rostral field (R) as previously described. The representation of high to low frequency tones in A1 was largely caudorostral along the plane of the sulcus. A reversal of the order of representation of frequencies occurred in R. (3) AI and R together were coextensive with a koniocellular, densely myelinated zone that expressed high levels of AchE and CO. These architectonic features were somewhat less pronounced in R than AI, but a clear border between the two areas was not apparent. (4) Cortex bordering AI and R was less responsive to tones, but when best frequencies for neurons could be determined, they matched those for adjoining parts of AI and R. (5) Architectonically distinct regions were apparent within some of the cortex bordering AI and R. (6) The major ipsilateral cortical connections of AI were with R and cortex immediately lateral and medial to AI. (7) Callosal connections of AI were predominantly with matched locations in the opposite AI, but they also included adjoining fields. (8) Neurons in the ventral (MGV), medial (MGM), and dorsal (MGD) nuclei of the medial geniculate complex projected to AI and cortex lateral to AI. (9) Injections in cortex responsive to high frequency tones labeled more dorsal parts of MGV than injections in cortex responsive to low frequency tones.

Acetylcholinesterase↗
PubMed1997

[Effects of salicylate and quinine on cat primary auditory cortex--spontaneous firing rate].

The effect of salicylate and quinine on the spontaneous firing rate in the cat primary auditory cortex was investigated in 13 healthy cats. Spontaneous firing rates were calculated for each single unit. A dose of 200 mg of sodium salicylate per kg was administered intraperitoneally to six cats, and the findings from the same single unit were recorded prior to application and continuously up to, on average, 6 hours after application. A dose of 100 or 200 mg of quinine hydrochloride per kg was administered intramuscularly to seven cats, and the findings from the same single unit were recorded in the same manner as for the cats treated with sodium salicylate. Twenty one single units in salicylate-treated cats and 29 single units in quinine-treated cats were evaluated. All animals treated with salicylate showed a 20-30 dB threshold shift about 2 hours after application and showed no recovery during the course of the investigation. All animals treated with quinine showed a 10-40 dB threshold shift about 1 hour after application and recovered during the course of the investigation. There was no consistent difference in overall spontaneous firing rate before and after application in either salicylate-treated cats or quinine treated cats. In order to investigate a potential different effect on units with different spontaneous firing rates, we divided the cats into two groups, a high-firing rate group (pre-application firing rate > 1 spike/s) and a low firing rate group (pre-application firing rate < 1 spike/s). A significant decrease in the high-firing rate group (p < 0.05) and a significant increase in the low firing rate (p < 0.01) were observed in salicylate-treated cats. The same tendencies were observed in quinine-treated cats, but only the difference in the low firing rate group was significant (p < 0.05). The difference in the high-firing rate group was close to the significant level (p = 0.055). These changes in spontaneous firing rates in cat primary auditory cortex may be related to the generation of tinnitus.

Animals↗
PubMed1993

Differential effects of age on click-rate and amplitude modulation-frequency coding in primary auditory cortex of the cat.

Recordings were made from 185 neurons in the primary auditory cortex of cats in the age range of 15 to 297 days. A comparison was made between the tuning for click repetition-rate and for amplitude modulation-frequency of a noise burst on the basis of temporal Modulation Transfer Functions (tMTF). 90 of the 185 units had a clear band-pass type tMTF for both repetition rate and modulation frequency, there was, however, no correlation between the respective Best Modulation Frequencies (BMF). Amplitude modulated noise (AMnoise) was the more effective stimulus in young kittens while click-train stimulation was more effective in adult cats. For all neurons with significant synchronization, BMFs for both click-train and AMnoise increased with age from about 4 Hz in kittens younger than 30 days to about 10 Hz in adult cats. In the approximately 50% of the neurons that were tuned both to click rate and modulation frequency, however, the BMF to AMnoise was consistently and significantly higher than that for clicks. In this group the mean BMF for kittens younger than 30 days were 7.94 Hz for AMnoise and 3.29 Hz for clicks and in the adults 10.91 Hz for AMnoise and 7.71 Hz for clicks.

Acoustic Stimulation↗
PubMed2006

Development of contralateral and ipsilateral frequency representations in ferret primary auditory cortex.

Little is known about the maturation of functional maps in the primary auditory cortex (A1) after the onset of sensory experience. We used intrinsic signal imaging to examine the development of the tonotopic organization of ferret A1 with respect to contralateral and ipsilateral tone stimulation. Sound-evoked responses were recorded as early as postnatal day (P) 33, a few days after hearing onset. From P36 onwards, pure tone stimuli evoked restricted, tonotopically organized patches of activity. There was an age-dependent increase in the cortical area representing each octave, with a disproportionate expansion of cortical territory representing frequencies > 4 kHz after P60. Similar tonotopic maps were observed following stimulation of the contralateral and ipsilateral ears. During the first few weeks following hearing onset, no differences were found in the area of cortical activation or in the magnitude of the optical responses evoked by stimulation of each ear. In older animals, however, contralateral stimuli evoked stronger responses and activated a larger A1 area than ipsilateral stimuli. Our findings indicate that neither the tonotopic organization nor the representation of inputs from each ear reach maturity until approximately 1 month after hearing onset. These results have important implications for cortical signal processing in juvenile animals.

Acoustic Stimulation↗
PubMed1995

Synaptic interactions involving acetylcholine, glutamate, and GABA in rat auditory cortex.

Using electrophysiological techniques in the in vitro rat auditory cortex, we have examined how spontaneous acetylcholine (ACh) release modifies synaptic potentials mediated by glutamate and gamma-aminobutyric acid (GABA). Single stimulus pulses to lower layer VI elicited in layer III a four-component (A-D) extracellular field response involving synaptic potentials mediated by glutamate and GABA. The cholinesterases inhibitor eserine (10-20 microM) or the cholinergic agonist carbachol (25-50 microM) depressed by 10-50% the glutamatergic components A and C, and the GABAergic components B and D. Atropine reversed the depressive effects of eserine and carbachol. A novel finding was that the degree of depression of component A varied inversely with stimulus intensity. However, during partial pharmacological antagonism of GABAA receptors, depression of A varied directly, not inversely, with stimulus intensity. Normally, then, depression of A is offset by reduced GABAergic inhibition of A. We also tested for differential depression of responses mediated by N-methyl-D-aspartate (NMDA) versus non-NMDA glutamate receptors. Following physiological and pharmacological isolation of the responses, eserine depressed the non-NMDA, but not the NMDA, receptor-mediated potential. Since the isolated NMDA potential still could be depressed by carbachol, the data suggested that activation of NMDA receptors may reduce spontaneous ACh release. In support of this, preincubation of slices in NMDA (10-20 microM) largely prevented eserine's, but not carbachol's, depression of components A and B. These results permit three conclusions of relevance to cortical information processing: (1) spontaneous ACh release tonically depresses synaptic potentials mediated by glutamate and GABA; (2) ACh depresses responses to weak inputs to a greater degree than responses to strong inputs: (3) activation of NMDA receptors may "feedback" to reduce ACh release, a mechanism that could place regulation of local ACh release under glutamatergic afferent control.

Acetylcholine↗
PubMed1978

Single-unit activity in the auditory cortex of monkeys selectively attending left vs. right ear stimuli.

Single-unit responses were recorded from the auditory cortex of rhesus monkeys that were performing an auditory selective attention task. Acoustic stimuli were presented randomly to either the left or right ear through headphones. In a given block of trials one ear was selected as the ear to be attended, and was indicated by lighting either a left or right response key. The animal's task was to press the lighted key whenever stimuli were presented to the attended ear, but to make no response to stimuli presented to the other ear. The attended ear was alternated on successive blocks of trials. Fourteen of 77 units showed significantly greater rates of evoked discharges for an attended stimulus than for an identical non-attended stimulus. The increase in stimulus-evoked activity was not accompanied by any increase in spontaneous activity or by any changes in the pattern of a unit's response. Changes in firing rate occurred at latencies as early as 20 msec.

Acoustic Stimulation↗
PubMed1985

Patterns of organization in auditory cortex.

Previous work has described recently developed techniques that are contributing to advances in the study of the auditory system; some of these techniques have been used to make significant progress in this field. This paper includes a discussion of the tonotopic and binaural organizations of the auditory cortex (especially area AI), the neuronal connections between auditory cortical fields, the relationships between the functional maps and the connectivity patterns, the nature of binaural processing within AI, and the techniques used to assess the structural and functional properties of this portion of the central auditory pathway.

Animals↗