Receptive fields of auditory cortical neurons in the cat

Receptive fields of auditory cortical neurons were studied by electrical stimulation of nerve fibers in different parts of the cochlea in cats anesthetized with pentobarbital. The dimensions of the receptive fields were shown to depend on the topographic arrangement of the neuron in the auditory cortex. The more caudad the neuron on the cortical projection of the cochlea in the primary auditory cortex, the more extensive its receptive field. The receptive fields were narrowest in the basal turn of the cochlea and were symmetrical with respect to their center. It is suggested that the region of finest discrimination of acoustic stimuli in cats is located in the basal region of the cochlea, i.e., in that part of its receptor system which has the narrowest receptive field and is represented by significantly more (than the middle and apical regions of the cochlea) nerve cells in the primary auditory cortex [1].A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 13, No. 5, pp. 467–473, September–October, 1981.     

Cochleotopic organization of the second auditory cortical area in the cat

The cochleotopic organization of the second auditory cortical area was investigated in cats anesthetized with pentobarbital by the evoked potentials method. Two independent representations of the cochlea were shown to exist in area AII: One in the dorsocaudal portion, the other in its ventrorostral portion. These projections of the cochlea differ in size and in the order of representation of its different parts. The dorsocaudal part of the auditory projection area of the cochlea, which extends over a distance of 2.6–2.8 mm from the center of the basal to the center of the apical focus, is arc-shaped. The order of arrangement of projections of different parts of the cochlea in this region of the auditory cortex coincides with that in the first auditory area, whereas the projection of the cochlea in the ventrorostral part of area AII, the length of which is 1.4–1.6 mm, has the opposite order of representation. The localization of projections of the cochlea in different cats shows considerable variability not only as regards anatomical topography of the auditory cortex, but also from one animal to another. The basal region of the cochlea was shown to project to a larger area of the cortex than the middle and apical portions taken together. It is suggested that the basal turn of the cochlea is functionally the most important for perception and primary analysis of auditory information.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 1, pp. 18–27, January–February, 1980.     

Unit responses in the cat auditory cortex to electrical stimulation of nerve fibers innervating receptor cells in different parts of the organ of corti

Responses of 200 primary auditory cortical neurons to electrical stimulation of nerve fibers in different receptor zones of the cochlea were studied in cats anesthetized with pentobarbital. Under the influence of paired stimulation, after the response to the conditioning stimulus a state of prolonged (from 4 to 200 msec) refractiveness to the second stimulus developed in all the neurons tested. This long-lasting inhibition of unit activity was due to inhibition developing in the thalamus and the auditory cortex itself. The intensity and duration of excitation and inhibition in the cortical projection focus were maximal when the center of the receptive field was stimulated and decreased when the stimulus shifted from the center to the periphery. The region of the receptor surface of the cochlea to stimulation of which the auditory cortical neurons respond by an action potential is much narrower than the region whose electrical stimulation depresses the discharge of these neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 14, No. 4, pp. 418–425, July–August, 1982.     

Association connections of the cat's parietal cortex

Intercortical connections of primary sensory (visual, auditory, somatosensory) areas with the parietal association cortex were studied in cats by the retrograde axonal transport of horseradish peroxidase and the Fink-Heimer silver impregnation of degenerated fibers techniques. This combined study revealed the shape, size, and intracortical location of cells connecting the primary sensory areas monosynaptically with the parietal cortex and also the distribution of preterminals and terminals of the fibers of these cells in the parietal association cortex. The greatest number of cells forming connections with area 7 of the parietal association cortex was shown to occur in visual area V1, and with area 5 in somatosensory area S1. Besides pyramidal neurons tagged with horseradish peroxidase, which were located mainly in layers II–IV, a few tagged stellate and fusiform cells also were found. The results supplement and confirm data on afferent connections of the parietal association cortex in cats.M. Gor'kii Donetsk Medical Institute. Translated from Neirofiziologiya, Vol. 13, No. 1, pp. 3–6, January, 1981.     

Comparison of spontaneous cortical unit activity in several brain areas of unanesthetized cats

Spontaneous unit activity in association area 5 and some projection areas of the cortex (first somatosensory, first and second auditory areas) were studied in cats immobilized with D-tubocurarine in which the index of specific spontaneous activity, the mean frequency, types of spontaneous activity, and statistical parameters — distribution of interspike intervals and autocorrelation function — were determined. The results showed that spontaneous unit activity in the association area differs from that in the projection areas in both intensity and character. A special feature of the spontaneous activity of the auditory areas was a well-marked volley distribution of activity. In the somatosensory area the level of spontaneous activity as reflected in all indices was the lowest. In the association cortex the largest number of neurons with spontaneous activity lay at a depth of 500–1000 µ corresponding to cortical layers III–IV. In the first auditory area neurons with spontaneous activity were concentrated at a depth of 1400 µ (layer V) and in the somatosensory area at a depth of 1000–1400 µ (alyers IV–V). The possible functional significance of these differences is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 1, pp. 13–21, January–February, 1976.     

Morphological characteristics of the neuronal population in the feline auditory cortex projecting into the medial geniculate body

The location and morphological profile of auditory cortex neurons projecting to the medial geniculate body were investigated in adult cats using horseradish peroxidase retrograde axonal transport techniques. Sources of descending projections to the medial geniculate body from auditory cortex areas I and II were found to be neurons belonging to deep-lying layers (layer VI and layer V to a lesser extent). By far the majority of corticogeniculate neurons in the auditory cortex were pyramidal cells. In layer VI of the primary auditory area (A1), the number of corticogeniculate neurons reaches 60% of all cells belonging to that layer. The average area (M±m) of the profile of perikarya of corticogeniculate neurons in layer VI, area Al equaled 139.3±2.5 µm² and 219.5±7.0 µm² in layer V neurons; average size of long diameter: 15.0±0.19 and 18.3±0.4 µm respectively. The lower regions of layers III and IV in area Al were found to be the termination point of the greater mass of anterogradely-labeled geniculocortical fibers (terminals of relay neuron axons belonging to the medial geniculate body).A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 4, July–August, pp. 513–521, 1989.     

Single unit activity related to acoustic stimulus meaning in the cat auditory cortex during instrumental food reflex

Responses of 93 neurons to isolated presentation of a single click and a series of 10 clicks with following frequency of 1000 Hz and responses of 66 neurons after the click had become a positive conditioned stimulus, and a series of 10 clicks had become a differential, negative stimulus, were investigated in chronic experiments on cats. Formation and realization of differential inhibition of an instrumental food reflex was shown not to lead to strengthening of inhibition in the auditory cortex, and the process of differential inhibition itself within the primary auditory cortex is not essentially an inhibitory process. Identical changes were found in responses of auditory cortical neurons to both positive and negative conditioned stimuli after training, evidence that neurons of the primary auditory cortex play a similar role in realization of the instrumental reflex and in its differential inhibition. It is suggested that the presence of groups of neurons responding by excitation or inhibition only to presentation of a stimulus with definite informative value is of great importance for differentiation.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukranian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 2, pp. 212–221, March–April, 1985.     

Effect of pentobarbital on neuronal inhibitory response evoked in an isolated slab of cat auditory cortex by intracortical stimulation

Neuronal responses of an acutely isolated slab of auditory cortex (area AI) to intracortical electrical stimulation were studied intracellularly in cats anesthetized with pentobarbital. It was found that 77% of responses were primary IPSPs, and allowing for secondary inhibitory responses, an inhibitory response was observed in 92% of neurons. All types of neuronal responses in the slab were short-latency. The maximal response latency did not exceed 5 msec. Neurons responding to stimulation by IPSPs were found at all depths in the slab, with a maximum in layers II–III. Nearly all primary IPSPswere mono- and disynaptic. Pentobarbital increased the duration of individual neuronal inhibitory responses in the isolated slab of auditory cortex without affecting maximal duration of the IPSP. The mechanisms of the effect of pentobarbital on the amplitude and duration of IPSPs are discussed.I. I. Mechnikov Odessa State University. Translated from Neirofiziologiya, Vol. 16, No. 2, pp. 147–152, March–April, 1984.     

Changes in amplitude and temporal characteristics of evoked potentials in the sensomotor cortex after division of the spinocervical tracts in cats

Temporal and amplitude characteristics of evoked potentials of the sensomotor cortex in waking cats were studied during variation in the intensity of electrodermal stimulation. The results obtained in experiments on intact animals and on the same animals for several months after division of the spinocervical tracts at the cervical level were compared. After blocking of the inflow of afferent impulses along these tracts of the spinal cord, statistically significant changes in evoked potentials were observed, mainly in response to medium and strong stimulation. These changes were more clear in the motor and second somatosensory areas of the cortex. A decrease in sensitivity to pain also was found. During recovery of the motor functions, cutaneous sensation remained impaired and the amplitude characteristics of the evoked somatosensory activity were not restored. The results suggest that thinner fibers predominate among the primary afferent fibers of the spinocervical tract, and their projections are more widely represented in the second somatosensory and motor areas of the cortex.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 4, No. 5, pp. 516–523, September–October, 1972.     

Morphology of the cat auditory cortex

The ultrastructural features of the primary auditory cortex of the cats and the character of the endings of geniculo-cortical afferent fibers in the early stages of experimental degeneration evoked by destruction of the medial geniculate body were studied. In all layers of the cortex asymmetrical synapses with round synaptic vesicles on dendritic spines and on thin dendritic branches of pyramidal and nonpyramidal neurons are predominant. Symmetrical synapses with flattened or polymorphic vesicles are distributed chiefly on the bodies of the neurons and their large dendrites. Because there are few symmetrical synapses which could be regarded as inhibitory it is postulated that inhibitory influences may also be transmitted through asymmetrical synapses with round vesicles. Other types of contacts between the bodies of neurons, dendrites, and glial processes also were found in the auditory cortex. Degenerating terminals of geniculo-cortical fibers were shown to terminate chiefly in layer IV of the cortex on pyramidal and nonpyramidal neurons. Degeneration was of the dark type in asymmetrical synapses with round vesicles. The results are dicussed in connection with electrophysiological investigations of the auditory cortex.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 5, pp. 519–524, September–October, 1973.     

Regional variations in spontaneous unit activity in the neuronally isolated cortex

Unit activity of the visual, auditory, and association areas of the neuronally isolated cortex of one hemisphere was studied in acute experiments on cats (the method was described previously [5]). A few hours after operation, the same types of spontaneous unit activity are observed in the isolated cortex as in the intact cortex under identical experimental conditions. Unlike in the normal cortex, most cells in the neuronally isolated cortex discharge regularly. Spontaneously active cells are two or three times fewer in the isolated cortex than in the intact. The distribution of active cells by depth in the neuronally isolated cortex varies from one region to another.Institute of Experimental Medicine, Academy of Medical Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 2, No. 5, pp. 475–481, September–October, 1970.     

Area 39 and 41 neurons sensitive to acoustic stimulation in the rat cortex: Polysensory properties

A comparative analysis of the polysensory properties of 102 neurons in areas 39 and 41 (the associative and auditory cortices, respectively) was performed in acute experiments on rats under chloralose-nembutal anesthesia. In the auditory cortex, the evoked potentials (EP) recorded from the surface of the above area in response to acoustic tonal, electrical cutaneous, and light stimulations almost always were distinguished by their shorter (4–5 msec) latency and higher amplitude. We studied neurons in both areas; their responses to the pure tones of various frequencies and to the stimulations of other modalities were compared. Bi- and polysensory neurons constituted 56.4% in area 39, and only 23% in area 41. The depth distribution of the responding neurons in areas 39 and 41 was different. Neurons with selective sensitivity to different frequencies of tonal signals were found in both areas. Usually monomodal neurons demonstrated selective properties in the auditory cortex, and 70% of them had a characteristic frequency. Over one-half of polymodal cells were frequency-selective in the associative cortex.Neirofiziologiya/Neurophysiology, Vol. 26, No. 3, pp. 223–229, May–June, 1994.     

Unit responses in the auditory cortex to stimulation of geniculocortical fibers

Extracellular and intracellular single unit responses of neurons of the auditory cortex to electrical stimulation of geniculocortical fibers (GCF) were recorded in experiments on cats immobilized with tubocurarine. The latent period of responses of 15% of neurons to GCF stimulation was 0.3–1.5 msec. It is postulated that they were excited anti-dromically. The latent period of spikes generated by neurons responding to GCF stimulation orthodromically varied from 1.6 to 12 msec. In 28.6% of neurons the latent period was 1.6–2.5 msec. It is postulated that these neurons were excited monosynaptically. Intracellular recording revealed primary IPSPs in response to GCF stimulation in 63.3% of neurons, a brief EPSP followed by a prolonged IPSP in 17.7%, an EPSP-spike-IPSP complex in 12.3%, and subthreshold EPSPs in 7% of neurons. The latent period of the primary IPSPs varied from 1.8 to 11 msec, being 1.8–3.7 in 72%, 3.8–5.7 in 20.0%, and 5.8–11 msec in 8.0% of neurons. The latent period of responses beginning with an EPSP was 1–4 msec (mean 1.8 msec). Orthodromic responses arising 3–10 msec after the antidromic response, and consisting of 3–5 spikes, were recorded in some antidromically excited neurons. Hypotheses regarding the functional organization of the auditory cortex and mechanisms of inhibition in its neurons are put forward on the basis of the results obtained.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 3, pp. 227–235, May–June, 1972.     

Unit responses in the cat auditory cortex to medial geniculate body stimulation

Responses of 98 auditory cortical neurons to electrical stimulation of the medial geniculate body (MGB) were recorded (45 extracellulary, 53 intracellularly) in experiments on cats immobilized with tubocurarine. Responses of the same neurons to clicks were recorded for comparison. Of the total number of neurons, 75 (76%) responded both to MGB stimulation and to clicks, and 23 (24%) to MGB stimulation only. The latent period of extracellularly recorded action potentials of auditory cortical neurons in response to clicks varied from 7 to 28 msec (late responses were disregarded), and that to MGB stimulation varied from 1.5 to 12.5 msec. For EPSPs these values were 8–13 and 1–4 msec respectively. The latent period of IPSPs arising in response to MGB stimulation varied from 2.2 to 6.5 msec; for 34% of neurons it did not exceed 3 msec. The difference between the latent periods of responses to clicks and to MGB stimulation varied for different neurons from 6 to 21 msec. Responses of 11% of neurons to MGB stimulation, recorded intracellularly, consisted of sub-threshold EPSPs, while responses of 23% of neurons began with an EPSP which was either followed by an action potential and subsequent IPSP or was at once cut off by an IPSP; 66% of neurons responded with primary IPSPs. Neurons responding to MGB stimulation by primary IPSPs are distributed irregularly in the depth of the cortex: there are very few in layers III and IV and many more at a depth of 1.6–2 mm. Conversely, excited neurons are predominant in layer III and IV, and they are few in number at a depth of 1.6–2 mm. It is concluded that the afferent volley reaching the auditory cortex induces excitation of some neurons therein and, at the same time, by the principle of reciprocity, induces inhibition of others. This afferent inhibition takes place with the participation of inhibitory interneurons, and in some cells the inhibition is recurrent. The existence of reciprocal relationships between neurons in different layers of the auditory cortex is postulated.A. A. Bogomolets' Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 1, pp. 23–31, January–February, 1972.     

Monosynaptic inhibitory postsynaptic potentials of cortical neurons

Monopolar intracortical stimulation of the auditory cortex was carried out in cats immobilized with D-tubocurarine. A macroelectrode (tip diameter 100 µ) or a microelectrode (tip diameter 10–15 µ) was used for stimulation. In both cases, besides excitatory responses, primary IPSPs with latent periods of 0.4–1.2 and 1.4–6.0 msec were recorded in cortical neurons close to the point of stimulation. The first group of IPSPs are considered to be generated in response to direct stimulation of bodies or axons of inhibitory cortical neurons, i.e., monosynaptically. The amplitude of these IPSPs varied in different neurons from 3 to 15 mV, and their duration from 4 to 150 msec. Additional later inhibitory responses were superposed on many of them. Of the IPSPs generated in auditory cortical neurons in response to stimulation of geniculocortical fibers 1.5% had a latency of 0.8–1.3 msec. They also are assumed to be monosynaptic. It is concluded that the duration of synaptic delay of IPSPs in cortical neurons and spinal motoneurons is the same, namely 0.3–0.4 msec. Axons of auditory cortical inhibitory neurons may be 1.5 mm long. The velocity of impulse conduction along these axons is 1.6–2.8 m/sec. The genesis of some special features of IPSPs of cortical neurons is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 7, No. 5, pp. 458–467, September–October, 1975.     

Responses of parietal associative neurons to stimulation of primary sensory areas

Responses of 251 neurons in the anterior part of the middle suprasylvian gyrus to stimulation of primary sensory (auditory, visual, somatosensory) areas and also to acoustic, visual, and somatosensory stimuli were studied in acute experiments on cats anesthetized with chloralose (40 mg/kg) and pentobarbital (20 mg/kg). Three groups of neurons were distinguished by their responses to stimulation of the primary sensory areas: those responding by an increased firing rate (117) or by inhibition (35) and those not responding (99). Responses of 193 neurons to stimulation of the peripheral afferent systems were analyzed. Neurons of the parietal associative cortex responded more frequently to cortical stimulation than to peripheral. By the duration of the latent period of their response to cortical stimulation the neurons were divided into three groups: those with short (less than 20 msec), medium (20–30 msec), and long latent periods (over 30 msec). The first group was the largest.Kemerovo State Medical Institute. Translated from Neirofiziologiya, Vol. 4, No. 5, pp. 524–530, September–October, 1972.     

Responses of medial geniculate body neurons to auditory cortical stimulation

Extracellular and intracellular unit responses of thepars principalis of the medial geniculate body to stimulation of the first (AI), second (AII), and third (AIII) auditory cortical areas were studied in cats immobilized with D-tubocurarine. In response to auditory cortical stimulation both antidromic (45–50%) and orthodromic (50–55%) responses occurred in the geniculate neurons. The latent period of the antidromic responses was 0.3–2.5 msec and of the orthodromic 2.0–18.0 msec. Late responses had a latent period of 30–200 msec. Of all neurons responding antidromically to stimulation of AII, 63% responded antidromically to stimulation of AI also, confirming the hypothesis that many of the same neurons of the medial geniculate body have projections into both auditory areas. Orthodromic responses of geniculate neurons consisted either of 1 or 2 spikes or of volleys of 8–12 spikes with a frequency of 300–600/sec. It is suggested that the volleys of spikes were discharges of inhibitory neurons. Intracellular responses were recorded in the form of antidromic spikes, EPSPs, EPSP-spike, EPSP-spike-IPSP, EPSP-IPSP, and primary IPSP. Over 50% of primary IPSP had a latent period of 2.0–4.0 msec. It is suggested that they arose through the participation of inhibitory interneurons located in the medial geniculate body.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 1, pp. 5–12, January–February, 1976.     

Unit responses of the cat caudate nucleus to cortical stimulation before and after destruction of nonspecific thalamic nuclei

Responses of caudate neurons to stimulation of the anterior sigmoid and various parts of the suprasylvian gyrus were studied in acute experiments on cats. The experiments consisted of two series: on animals with an intact thalamus and on animals after preliminary destruction of the nonspecific thalamic nuclei. Stimulation of all cortical areas tested in intact animals evoked complex multicomponent responses in caudate neurons with (or without) initial excitation, followed by a phase of inhibition and late activation. The latent periods of the initial responses to stimulation of all parts of the cortex were long and averaged 14.5–25.5 msec. Quantitative and qualitative differences were established in responses of the caudate neurons to stimulation of different parts of the cortex. Considerable convergence of cortical influences on neurons of the caudate nucleus was found. After destruction of the nonspecific thalamic nuclei all components of the complex response of the caudate neurons to cortical stimulation were preserved, and only the time course of late activation was modified.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 5, pp. 464–471, September–October, 1980.     

Electrophysiological analysis of the functional organization of cortico-cerebellar connections

The effect of stimulation of cortical association (orbito-frontal, parietal) and projection (auditory, sensomotor) areas on the activity of Purkinje neurons of the cerebellar cortex was studied in adult cats anesthetized with pentobarbital, with or without chloralose. These responses were compared with those to peripheral stimuli. Definite similarity was found between the responses of Purkinje cells to different cortical (association and projection) stimuli as regards both the types of responses of the neurons and their ability to respond. No similarity was observed in the responses of Purkinje cells to peripheral (visual, auditory, electrodermal) stimulation. Whereas almost identical numbers of neurons (over 50%) were excited in response to the different forms of cortical stimulation, the ability of the neurons to respond to peripheral stimuli differed considerably: 44.6% of neurons responded to electrodermal stimulation, 34.2% to auditory, and 18.8% to visual.Medical Institute, Kemerovo. Translated from Neirofiziologiya, Vol. 8, No. 5, pp. 483–489, September–October, 1976.     

Response of cortical evoked potentials to electric stimulation of the acoustic nerve in animals with damaged cochlea

Evoked potentials of the auditory cortex during the electrical stimulation of the cochlea were studied in acute experiments on cats. A series of electric pulses of short duration and different frequency delivered to the streptomycin-damaged cochlea were used as a stimulus. It has been shown that an amplitude and latency of electrical cortex responses depended on the number of pulses in series and on the interpulse intervals. Amplitudes of evoked responses increased with the growth of the number of stimuli. Latent periods changed in a narrower stimulation frequency band. Dependence of the induced potentials' amplitude growth on the increase in the number of electric pulses changed as a result of the two-fold enhancement of the stimulation amplitude.