Areas Of Auditory Cortex Research Articles

Cortical projections to the superior colliculus in the macaque monkey: a retrograde study using horseradish peroxidase.

The topical and laminar distribution of corticotectal cells, as well as their size and morphology, were studied in the macaque monkey with the horseradish peroxidase (HRP) technique. After HRP injections restricted primarily to the superficial layers of the colliculus, labelled cells were found in visual cortex (areas 17, 18, and 19) and both in the frontal eye field (area 8) and the adjacent part of premotor cortex (area 6). The clustering of labelled cells in visual cortex indicated that each of the anatomically and functionally distinct visual areas has its own set of collicular projections. When intermediate and deeper layers of the colliculus were injected, labelled cells were found also in posterior parietal cortex (area 7) where they were concentrated mainly on the posterior bank of the intraparietal fissure, in inferotemporal cortex (areas 20 and 21), in auditory cortex (area 22), in the somatosensory representation SII (anterior bank of sylvian fissure, area 2), in upper insular cortex (area 14), in motor cortex (area 4), in premotor cortex (area 6), and in prefrontal cortex (area 9). In the motor and premotor cortex, labelled cells formed a continuous band which appeared to stretch across finger-hand-arm-shoulder-neck representation. Similarly, the cluster of labelled cells in area 2 may correspond to the finger-hand representation of SII. The cortical regions not containing labelled cells were the somatosensory representation SI (areas 3, 1 and 2) and the infraorbital cortex. Labelled cells were restricted to layer V of all cortical areas except in the primary visual cortex, where labelled cells were found in both layer V and layer VI. The size spectrum of corticotectal cells ranged from 14.8 micron (average diameter) in area 17 to 27.8 micron in area 6, comprising cells as small as 8 micron and as large as 45 micron. Labelled cells in posterior parietal (area 7), in auditory (area 22), and in motor cortex (area 4) were small and distributed over only a narrow range of sizes. Those in premotor cortex (area 6) were often large and had a wide range in size distribution. The differences in size and morphology of corticotectal neurons suggest that they do not form a uniform class of neurons.

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.

Intrinsic organization of the cat's medial geniculate body identified by projections to binaural response-specific bands in the primary auditory cortex

The area of the cat's primary auditory cortex (AI) within which high frequency sounds are represented can be subdivided using functional criteria. Within each subdivision, or "binaural interaction band," all recorded neurons display similar responses to binaural stimulation. The current study distinguishes the thalamic sources of input to these subdivisions of AI and characterizes the topography within the thalamic projection to each class of bands. The borders of binaural bands in AI were mapped using microelectrode recording with diotic tonal stimulation, then injections of one to three retrograde tracers were introduced into identified bands. Within the ventral division (V) of the medial geniculate body (the major thalamic source of input to AI), the neuronal populations that projected to different classes of binaural bands were strictly segregated from each other. This segregation of class-specific thalamic sources constitutes a laminar organization within an axis of V that is orthogonal to the previously described tonotopic organization. Excitatory/excitatory (EE) binaural neurons in AI were found to be segregated from excitatory/inhibitory (EI) neurons in alternating "bands." We consistently identified: (1) a ventral pair of rostrocaudally continuous EI and EE bands, (2) a middle area within which the pattern of binaural subdivisions was more variable and within which bands often were discontinuous rostrocaudally, and (3) a dorsal zone (DZ) within which the responses of neurons differed in binaural properties and in frequency specificity from the response patterns that were characteristic of neurons elsewhere in AI. Each EI band apparently derived input that converged from three thickened laminae of cells in V that were oriented approximately horizontally. The most ventral of these laminae encompassed the ovoidal part of V (Vo), suggesting that EI bands are the only recipients in AI of a projection from Vo. All of the EE bands and DZ derived their input from a single continuous structure which included the dorsal two-thirds of the rostral pole of V and a horizontal lamina interposed between the two dorsalmost EI-projecting laminae. Restricted portions of the complex EE-projecting structure in V projected preferentially to particular EE subdivisions of AI. The V-to-AI thalamocortical topography exhibited a high degree of convergence and divergence within the projections to each cortical binaural band and within the projections to each class of bands. These observations indicate that the high frequency representation in AI and its principal thalamic source of input, the ventral division of the medial geniculate body, may be thought of as assemblies of spatially discrete, functionally distinguishable subunits. The significance of this intrinsic organization is discussed in regard to the requirements for analysis of sound stimuli.

Dynamics of auditory cortical unit responses of the waking cat during defensive conditioning to acoustic stimulation

In chronic experiments on cats unit responses of the primary auditory cortex (area 50) were studied by microelectrode recording during defensive conditioning to sound. During formation of the reflex biphasic responses with relatively short-latency (50–100 msec) and longer-latency (400–500 and 800–900 msec) activation predominated. Neighboring neurons, whose activity was recorded by the same microelectrode, also were involved more intensively in activity. Application of a differential stimulus in 70% of cases produced definite changes in unit activity, among which responses of activation type predominated. Analysis of the course of spike responses of the same neuron during the period of action of a large number of combinations and its comparison with the formation of the conditioned-reflex motor response revealed no direct correlation between these events.

Spontaneous activity of auditory cortical neurons of waking cats at rest and during defensive conditioning

The results of a computerized statistical analysis of 366 realizations of spontaneous spike activity of 181 neurons in the primary auditory cortex (area 50) of waking cats at rest and during defensive conditioning are described. In both situations the parameters of spontaneous activity of most neurons differed from those of a random flow. Conditioning led, on the one hand, to a stable increase in the frequency of spontaneous activity in intertrial periods and, on the other hand, judging from changes in the mean firing rate, the coefficients of variation of the length of the interspike intervals, the histograms of their distribution, and also the increase in the number of neurons with different forms of correlation between interspike intervals, to an increase in its stability (degree of organization).

Aural representation in the Doppler-shifted-CF processing area of the auditory cortex of the mustache bat.

In the mustache bat (Pteronotus pamellii rubiginosus) the frequency and amplitude of an acoustic signal are represented in the coordinates parallel to the surface of the Doppler-shifted-CF (constant frequency) processing area ofthe primary auditory cortex. In this area all cortical neurons studied were excited by contralateral stimuli, and almost all of them were either excited or inhibited by ipsilateral stimuli. These are called E-E (ipsilateral and contralateral excitatory) and I-E (ipsilateral inhibitory and contralateral excitatory) neurons, respectively. The I-E neurons are directionally sensitive, while the E-E neurons are not. The E-E neurons are equally sensitive to echoes between 30 degrees contralateral and 30 degrees ipsilateral. Of the electrode penetrations orthogonal to the Doppler-shifted-CF processing area, 57 percent were characterized by either E-E or I-E neurons. Thus, there are at least two types of binaural columns: E-E columns, mainly located in a ventral part of the Doppler-shifted-CF processing area, where neurons are tuned to weak echoes; and IE columns, mainly distributed in a dorsal part, where neurons are tuned to moderate to intense echoes. Therefore, neurons tuned to weaker echoes integrate or even multiply faint signals from both ears for effective detection of a distant small target, while neurons tuned to moderate to intense echoes are suited for processing directional information and are stimulated when a bat approaches a target at short range. The Doppler-shifted-CF processing area may be considered to consist of two functional subdivisions.

Projections from the ‘Cingular’ vocalization area in the squirrel monkey

In 5 squirrel monkeys the anatomical projections from the 'cingular' vocalization area were studied by the autoradiographic tracing technique. The 'cingular' vocalization area lies around the sulcus cinguli at the level of the genu of the corpus callosum; its electrical stimulation yields purring and cackling calls. The following efferent connections were found: corticocortical fibers could be traced into the orbital cortex (areas 10 and 11), dorsomedial frontal cortex (areas 9, 8 and 6), limbic cortex (areas 25, 24 and 23), Broca's area (area 44), frontal operculum (area 50), insula (areas 13 and 14), and auditory association cortex (area 22). Subcortical terminal fields within the telencephalon were found in the nucleus caudatus, putamen, claustrum, globus pallidus, olfactory tubercle, preoptic region and nucleus centralis and basolateralis amygdalae. Fibers reached most of these structures along different trajectories. In the diencephalon terminal fields lay in the dorsal hypothalamus, the subthalamus, lateral habenular nucleus, and the following thalamic nuclei: nucleus reticularis, ventralis anterior, centralis medialis, centralis superior lateralis, centralis inferior, submedius, medialis dorsalis and centrum medianum. In the midbrain, the periaqueductal gray was the only projection area, extending into the parabrachial nuclei at the pontomesencephalic transition. The most caudal terminal field was found in the medial pontine gray. No terminals were detected in the nucleus ambiguus, nucleus n. hypoglossi or in any other cranial motor nucleus involved in phonation processes. A comparison of this projection system with the whole of structures producing vocalization when electrically stimulated yielded only partial overlap. Not all vocalization areas lie within the 'cingular' projection system, and inversely, not the whole projection system yielded vocalization. Overlap took place in the anterior limbic cortex, preoptic region, central amygdaloid nucleus, midline thalamus, dorsal hypothalamus, periaqueductal gray and parabrachial nuclei. These structures are considered to compose a functionally coherent vocalization system. The projections into Broca's area, nucleus ventralis anterior thalami, frontoopercular cortex within the lateral fissure, pontine nuclei and superior temporal gyrus are discussed in their possible relationship to vocalization processes.

Electric reactions of the auditory cortex areas in bats to ultrasonic stimuli with various fill frequencies

The overall electric reactions and action potentials of single neurons in the auditory cortex were investigated for Vespertilionidae (Myotis oxygnathus) and Rhinolophidae (Rhinolophus ferrum equinum) narcotized with Hexenal. In the Vespertilionidae the greatest sensitivity to ultrasound is manifest at frequencies from 10 to 50 kHz, and in the Rhinolophidae for the ranges from 10 to 40 and from 82 to 84 kHz. The shapes of the response areas of single neurons in both types of bats are similar except for neurons discovered in Rhinolophidae that have three response areas with characteristic frequencies in the ranges 27–28, 40–42, and 80–84kHz. Narrow response areas with characteristic frequencies in the range from 70 to 90kHz appear on a considerable proportion of the neurons in the Rhinolophidae, but not the Vespertilionidae. Low thresholds are recorded to the stimulus cutoff in the range from 76 to 86 kHz.