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Distribution of response types across entire hemispheres of the mustached bat's auditory cortex.

AILDD1 AC'1 The responses of neurons in the mustached bat's auditory cortex are specialized to extract particular information from biosonar signals. For this study, we mapped response properties across entire hemispheres in several animals. These experiments enabled us to construct a standard map that aided in determining the connections among the areas, as described subsequently. The mapping also yielded quantitative data regarding the relative sizes of areas and the proportion of cortex devoted to different response types. We identified six response types that were distributed in 11 areas. Eight areas, comprising two-thirds of the auditory cortex, contained neurons sensitive to particular components in biosonar signals. Most were facilitated by combinations of frequency modulated and constant frequency biosonar signal components (FMs and CFs, respectively). There were three major types of combination-sensitive neurons: FM-FM, CF/CF, and FM-CF. Each type of combination sensitivity occurred in multiple areas. The largest proportion were FM-FM neurons (approximately 30% of all neurons in auditory cortex), followed by FM1-CF2 (approximately 23%) and CF/CF (approximately 11%). In the other three areas comprising approximately one-third of the auditory cortex, most neurons responded well to frequencies not contained in biosonar signals.

Animals↗

Pyramidal cells in primary auditory cortex project to cochlear nucleus in rat.

Recent work has demonstrated that the auditory cortex in rat sends direct projections to the auditory nuclei of the brainstem, including the cochlear nucleus and superior olive. To determine the cortical origin of the projections to cochlear nucleus, Fast Blue, a retrograde fluorescent tracer, was injected into the cochlear nucleus. Labeled cells in the forebrain were then studied with light microscopy and mapped. The projection was found to originate from large pyramidal neurons in layer V of primary auditory cortex. The projection was predominantly ipsilateral, and no labeled neurons were found in other cortical areas. These data imply that primary auditory cortex exerts influence over ascending auditory information at the earliest stages of the central auditory system.

Amidines↗

A type of apparently axonless granule cell in the cat auditory cortex.

In the auditory areas (A I, A II and the posterior ectosylvian gyrus) of the cat, unusually small granule cells apparently lacking an axon are described. They are exclusively located in layer 2/3 and sometimes impregnate in groups of 4 or 5, being probably quite frequent. Some morphological differences between granule cells of young and adult cats are noted.

Aging↗

Topographic and temporal indices of vowel spectral envelope extraction in the human auditory cortex.

The auditory-evoked neuromagnetic field elicited by single vowel formants and two-formant vowels was recorded under active listening conditions using a 37-channel magnetometer. There were three single formants with formant frequencies of 200, 400, and 800 Hz, another single formant with a formant frequency of 2600 Hz, and three vowels that were constructed by linear superimposition of the high- onto one of the low-frequency formants. P50 m and N100 m latency values were inversely correlated with the formant frequency of single formants. A strong effect of formant frequency on source location was obtained along the postero-anterior axis, which is orthogonal to the well-established latero-medial tonotopic gradient. Regardless of whether single formants or first formants of vowels were considered, N100 m sources were more anterior and sustained field sources were more posterior for higher-frequency than for lower-frequency formants. The velocity of the apparent posterior-to-anterior movement across cortical surface of N100 m sources first reported by Rogers et al. [Rogers, R. L., Papanicolaou, A. C., Baumann, S. B., Saydjari, C., & Eisenberg, H. M. (1990). Neuromagnetic evidence of a dynamic excitation pattern generating the N100 auditory response. Electroencephalography and Clinical Neurophysiology,77, 237-240] decreased as a function of latency. The amount of deceleration was positively correlated with formant frequency. Responses to the vowels were superadditive, indicating that the processes elicited by the constituents of composite stimuli interact at one or more stages of the afferent auditory pathway. Such interaction may account for the absence of a lateral-to-medial tonotopic mapping of first formant frequency. The source topography found may reflect activity in auditory fields adjacent to AI with the strength of the contribution varying with formant frequency. Alternatively, it may reflect sharpness-of-tuning and inhibitory response-area asymmetry gradients along isofrequency stripes within AI. Either alternative may be interpreted in terms of a spectral blurring mechanism that abstracts spectral envelope information from the details of spectral composition, an important step towards the formation of invariant phonetic percepts.

Adult↗

Hierarchical organization of the human auditory cortex revealed by functional magnetic resonance imaging.

The concept of hierarchical processing--that the sensory world is broken down into basic features later integrated into more complex stimulus preferences--originated from investigations of the visual cortex. Recent studies of the auditory cortex in nonhuman primates revealed a comparable architecture, in which core areas, receiving direct input from the thalamus, in turn, provide input to a surrounding belt. Here functional magnetic resonance imaging (fMRI) shows that the human auditory cortex displays a similar hierarchical organization: pure tones (PTs) activate primarily the core, whereas belt areas prefer complex sounds, such as narrow-band noise bursts.

Adult↗

Arterial blood supply to the auditory cortex of the chinchilla.

Utilizing optical imaging we identified and named the arteries that supply the primary auditory cortex in the chinchilla (Chinchilla laniger). The primary auditory cortex is located 2-3 mm caudal to the medial cerebral artery and is supplied by it. Using corrosion casts and scanning electron microscopy we visualized the capillary networks in the auditory cortex and found regional variations in the densities of the capillary bed. We hypothesize that the uneven capillary densities observed in the auditory cortex correspond to neurologically more active areas.

Animals↗

Maturation of human auditory cortex: implications for speech perception.

This project traced the maturation of the human auditory cortex from midgestation to young adulthood, using immunostaining of axonal neurofilaments to determine the time of onset of rapid conduction. The study identified 3 developmental periods, each characterized by maturation of a different axonal system. During the perinatal period (3rd trimester to 4th postnatal month), neurofilament expression occurs only in axons of the marginal layer. These axons drive the structural and functional development of cells in the deeper cortical layers, but do not relay external stimuli. In early childhood (6 months to 5 years), maturing thalamocortical afferents to the deeper cortical layers are the first source of input to the auditory cortex from lower levels of the auditory system. During later childhood (5 to 12 years), maturation of commissural and association axons in the superficial cortical layers allows communication between different subdivisions of the auditory cortex, thus forming a basis for more complex cortical processing of auditory stimuli.

Adult↗

Effects of noradrenaline on frequency tuning of rat auditory cortex neurons.

The selectivity of rat auditory cortex neurons for pure tone frequency was studied during and after ionophoretic application (5-40 nA) of noradrenaline in urethane-anaesthetized rats. The dominant effect induced by noradrenaline was a significant decrease in spontaneous (93/268 cells) and evoked activity (133/268 cells) which outlasted the application. In the whole population of cells (n = 268) the signal-to-noise ratio, computed using as the signal either the mean evoked response or the response at the best frequency, was unchanged during noradrenaline application. It was significantly increased only for cells showing significantly decreased spontaneous activity, and was significantly decreased for cells showing increased spontaneous activity. Frequency selectivity was significantly increased for the whole population during and after noradrenaline application. It was also significantly increased for cells showing significantly decreased evoked activity, and was significantly decreased for cells showing increased evoked activity. The noradrenaline-induced inhibition was not blocked by propranolol (beta antagonist); it was blocked by prazosin (alpha1 antagonist) and partly mimicked by phenylephrine (alpha1 agonist). GABA, which also inhibited spontaneous and evoked activity, slightly increased the signal-to-noise ratio and significant increased frequency selectivity. However, when noradrenaline was ejected in the presence of bicuculline at doses that were able to block GABAergic inhibition, the inhibitory effects of noradrenaline on spontaneous and evoked activity were still observed. The possible function of noradrenaline-induced inhibitions in sensory cortices is briefly discussed.

Animals↗

[Modification of cochlear active micromechanism by auditory cortex].

OBJECTIVE: To investigate the modification of active micromechanism of outer hair cells by auditory cortex. METHODS: 1. Effects of selective attention on distortion product otoacoustic emissions (DPOAE) were analyzed in 9 human subjects with normal hearing. Visual tasks, two letters, O and Q, were alternatively displayed on a computer screen. Letter Q was target stimulus flashing randomly (probability was 20%), and letter O was non-target stimulus occurring regularly (probability was 80%). 2. In 13 guinea pigs, DPOAEs were measured before and after decerebration. RESULTS: 1. Visual tasks suppressed DPOAE significantly in both 1 kHz and 2 kHz (P < 0.05). 2. DPOAE reduced significantly in decerebrated guinea pigs. CONCLUSION: The study further demonstrates that the efferents from the auditory cortex do innervate the cochlea, and the efferent pathway passes from the auditory cortex, the super olive complex to the cochlea. Through this pathway the auditory cortex modulates the cochlear active micromechanism. The underlying mechanism was discussed.

Acoustic Impedance Tests↗

[Functional organization of the callosal connections in the auditory cortex of the cat].

Homotopic transcallosal potentials of the cat auditory cortex primary projection area (AI) had the greatest amplitude, the minimal temporal parameters and the maximal stability of characteristics as compared with the responses of the auditory cortex associative fields AII, AIV, Ep. Heterotopic transcallosal responses in the AI field occurred in stimulation of the field under the same name whereas in the Ep field the responses occurred both in stimulation of the same and the AI and AII fields of the opposite hemisphere. The structure of transcallosal connections of the auditory cortex primary projection field seems to be characterized by homotopy whereas in associative fields the role of heterotopic transcallosal interactions seems to increase. Such a structure of transcallosal connections seems to maintain the important role of interhemispheric interaction in mechanisms of spatial hearing.

Animals↗

Functional subregions in primary auditory cortex defined by thalamocortical terminal arbors: an electrophysiological and anterograde labeling study.

Several functional maps have been described in primary auditory cortex, including those related to frequency, tuning, latency, binaurality, and intensity. Many of these maps are arranged in a discontinuous or patchy manner. Similarly, thalamocortical projections arising from the ventral division of the medial geniculate body to the primary auditory cortex are also patchy. We used anterograde labeling and electrophysiological methods to examine the relationship between thalamocortical patches and auditory cortical maps. Biotinylated dextran-amine was deposited into physiologically characterized sites in the ventral division of the medial geniculate body of New Zealand white rabbits. Approximately 7 d later, the animal was again anesthetized and the ipsilateral auditory cortex was mapped with tungsten microelectrodes. Multi-unit physiological data were obtained for the following characteristics: best frequency (BF), binaurality, response type, latency, sharpness of tuning, and threshold. Immunocytochemical methods were used to reveal the injection site in the ventral division of the medial geniculate body as well as the anterogradely labeled thalamocortical afferents in the auditory cortex. In 86% of the cases (12 of 14), entry into a thalamocortical patch was associated with a marked change in physiological responses. A consistent BF and binaural class were usually observed within a patch. The patches appear to innervate distinct functional regions coding frequency and binaurality. A model is presented showing how patchy thalamocortical projections participate in the formation of tonotopic and binaural maps in primary auditory cortex.

Animals↗

Frequency representation in auditory cortex of the common marmoset (Callithrix jacchus jacchus).

The location and characteristics of the primary auditory cortex of the common marmoset, Callithrix jacchus jacchus, were determined in five anesthetized male adult animals by mapping the responses of cortical units and unit clusters to pure tone stimuli presented to the contralateral ear. The primary auditory cortex lies largely ventral to the lateral sulcus, the only major fissure on the lateral cortex of this smooth-brained primate, but in some animals it may extend significantly down the ventral bank of this sulcus. Responses are distributed such that low best frequencies are found rostroventrally whereas high best frequencies occur caudally. The disposition of frequency-band contours is fan-shaped, with contours separating low-frequency octaves nearly parallel to the lateral sulcus and high-frequency (greater than 8 kHz) contours perpendicular to that sulcus. Best frequencies range from 0.6 to 30 kHz across the primary field, but there is a disproportionate representation of the three octaves between 2 and 16 kHz. The most sensitive thresholds (as low as -2 dB SPL) are found between 7 and 9 kHz. The primary auditory cortex is similar in cytoarchitecture to that reported for the cat, showing a blurring of lamination in the middle layers (II-IV) and a preponderance of small cells in these merged layers, giving a highly granular appearance. The accessibility of the cochlear representation on the gyral surface makes the marmoset an attractive animal for studies of primate auditory cortex.

Animals↗

The termination of callosal fibres in the auditory cortex of the rat. A combined Golgi--electron microscope and degeneration study.

When the corpus callosum of the rat is sectioned, the callosal fibres in the cerebral cortex undergo degeneration. In the auditory cortex (area 41) the degenerating axon terminals form asymmetric synapses, and the vast majority of them synapse with dendritic spines. Some other synapse with the shafts of both spiny and smooth dendrites, and a few with the perikarya of non-pyramidal cells. The degenerating axon terminals are contained principally within layer II/III, in which they aggregate in patches. Using a technique in which neurons within the cortex are Golgi-impregnated, then gold-toned and examined in the electron microscope, it has been shown that the dendritic spines of pyramidal neurons with cell bodies in different layers receive the degenerating callosal afferents. The spines arise from the main apical dendritic shafts and their branches, from the dendrites of the apical tufts, and in some cases from the basal dendrites of the pyramidal neurons. The shafts of some pyramidal cell apical dendrites also form asymmetric synapses with callosal afferents. Since we have encountered no spiny non-pyramidal neurons in Golgi preparations of rat auditory cortex, and because other types of non-pyramidal cells have few dendritic spines, it is concluded that practically all of the dendritic spines synapsing with callosal afferents originate from pyramidal neurons.

Animals↗

Effects of changes in cortical arousal and of auditory cortex cooling on neuronal activity in the medial geniculate body.

The activity of cells in the medial geniculate body (MGB) of adult cats was recorded during different states of cortical arousal with and without cooling of the auditory cortex. In the absence of auditory cortex cooling, the overall mean unit spontaneous discharge rate was 49% higher during desynchroized Electrocorticogram (ECoG) periods (high cortical arousal). Responses to sound were somewhat more prominent vis-à-vis the spontaneous activity during periods of high arousal. Changes in spontaneous discharge rate associated with arousal shifts were significantly reduced during auditory cortex cooling. When the ECoG changed from desynchronized to synchronized activity, MGB cells showed a change in discharge pattern, typically characterized by an increase in both high-rate bursts and long-interval pauses. These changes were duplicated for most cells by cooling of the auditory cortex. Corticofugal fiber discharge thus has an effect on MGB neuronal activity which is dependent on the level of cortical arousal. This effect is most likely a result of direct corticogeniculate activity, though indirect auditory cortex - brainstem - MGB routes may also be involved.

Animals↗

Acetylcholine modifies neuronal acoustic rate-level functions in guinea pig auditory cortex by an action at muscarinic receptors.

Cholinergic modification of neuronal responsiveness in auditory cortex includes alteration of spontaneous and tone-evoked neuronal discharge. Previously it was suggested that the effects of acetylcholine (ACh) and muscarinic agonists on neuronal discharge resembled those due to increases in the intensity of acoustic stimuli (Ashe et al. 1989). To determine the relationship between neuronal modifications due to ACh acting at muscarinic receptors and those due to changes in stimulus intensity, we determined acoustic rate-level functions for neurons in the auditory cortex of barbiturate-anesthetized guinea pigs before, during and after administration of ACh. ACh facilitated acoustic rate-level functions in 82% of the cells tested. In addition, during ACh administration 66% of neurons responded to stimuli that were previously subthreshold, that is, ACh decreased the response threshold. Cholinergic facilitation of rate-level functions was attenuated by the general muscarinic antagonist atropine. The nature of the muscarinic receptors involved in the actions of ACh was further examined by presenting single tones before, during, and after administration of ACh and specific muscarinic receptor subtype antagonists, either pirenzepine (M1) or gallamine (M2). ACh-induced facilitation of spontaneous and tone evoked neuronal discharge was antagonized by pirenzepine, but not by gallamine, suggesting the involvement of the M1 muscarinic receptor subtype. These data indicate that ACh can facilitate stimulus-evoked responses and decrease response thresholds for neurons in auditory cortex, possibly via activation of M1 muscarinic receptors. Such effects of ACh acting at muscarinic receptors could underly cholinergic regulation of information processing in the auditory cortex.

Acetylcholine↗

Intrinsic optical signals from rat primary auditory cortex in response to sound stimuli presented to contralateral, ipsilateral and bilateral ears.

In the auditory cortex, primitive features of acoustic stimuli are represented for auditory scene analysis. A typical example of a feature representation is the tonotopic map, in which sound frequencies are spatially arranged in an orderly manner. Some neurons in the auditory cortex are sensitive to sound source location, which is another important feature for auditory scene analysis. In the present study, using the intrinsic optical imaging technique, we attempted to visualize the two-dimensional pattern of neuronal population responses in the primary auditory cortex of rats to pure tones presented at various frequencies and sound intensities. The observed arrangements of sound frequencies were consistent with those obtained by electrophysiological mapping, which indicates that our intrinsic optical recording can visualize populational responses of neurons. We also found different temporal patterns of intrinsic signals elicited in response to contralateral, ipsilateral, and bilateral ear stimulations. Finally we try to explain the observed differential time courses of intrinsic signal responses from the theoretical point of view on the conduction of neural activities, based on the so far anatomically identified neural pathways in the rodent auditory system.

Acoustic Stimulation↗

Contribution of auditory cortex to sound localization by the ferret (Mustela putorius).

Ferrets were tested in a semicircular apparatus to determine the effects of auditory cortical lesions on their ability to localize sounds in space. They were trained to initiate trials while facing forward in the apparatus, and sounds were presented from one of two loudspeakers located in the horizontal plane. Minimum audible angles were obtained for three different positions, viz., the left hemifield, with loudspeakers centered around -60 degrees azimuth; the right hemifield, with loudspeakers centered around +60 degrees azimuth; and the midline with loudspeakers centered around 0 degrees azimuth. Animals with large bilateral lesions had severe impairments in localizing a single click in the midline test. Following complete destruction of the auditory cortex performance was only marginally above the level expected by chance even at large angles of speaker separation. Severe impairments were also found in localization of single clicks in both left and right lateral fields. In contrast, bilateral lesions restricted to the primary auditory cortex resulted in minimal impairments in midline localization. The same lesions, however, produced severe impairments in localization of single clicks in both left and right lateral fields. Large unilateral lesions that destroyed auditory cortex in one hemisphere resulted in an inability to localize single clicks in the contralateral hemifield. In contrast, no impairments were found in the midline test or in the ipsilateral hemifield. Unilateral lesions of the primary auditory cortex resulted in severe contralateral field deficits equivalent to those seen following complete unilateral destruction of auditory cortex. No deficits were seen in either the midline or the ipsilateral tests.

Animals↗