Responses of auditory cortical neurons to paired clicks

In cats immobilized with tubocurarine, a paired-click method was used to determine the duration of the refractory period of 75 auditory cortical neurons responding to clicks with a latent period of up to 30 msec. Sixty-eight of the neurons exhibited no spontaneous activity, while in the other seven spontaneous activity was infrequent and irregular. It was found that a click makes responding neurons refractory to a second click for a long time. The duration of this refractory period is 3 to 700 msec; it is constant for each neuron, but varies from one neuron to another. A direct relationship was found between the number of neurons responding to the second click and the interval between the first and second clicks: the shorter the interval the fewer neurons respond to the second click. It is postulated that this dependence lies at the basis of the neurophysiological mechanism of perception and discrimination of short time intervals.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 3, pp. 227–235, May–June, 1970.     

Responses of neurons in the auditory cortex to sound stimuli

The evoked potential (EP) and the pulse activity of single auditory cortex neurons were recorded simultaneously in response to a click and to a tone for cats under nembutal and nembutal — chloralose anesthesia. Both extra- and intracellular taps were employed. The experiments showed that the reaction of auditory cortex neurons in response to a click lasts from 200 to 300 msec. It consists of pulse discharges from several groups of neurons. Out of 174 neurons observed 8 responded within 4 to 7 msec after a click (before the EP). One hundred and nine neurons reacted in the range from 7 to 25 msec which coincided with the initial electropositivity of the EP; 11 neurons were in the range from 40 to 100 msec and 4 were between 180 and 270 msec. Such a sequence of involvement of different neuron groups in the reaction is probably accounted for to a large extent by the time dispersion of the afferent volley. With an intracellular tap slow alterations of membrane potential were observed in the form of an EPSP with pulses together with subsequent hyperpolarization lasting 50 to 70 msec and slowly increasing depolarization that reached a maximum after 170 to 200 msec.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 1, No. 2, pp. 147–157, September–October, 1969.     

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.     

Responses of auditory cortex neurons in unanesthetized cats to best-frequency tones

The characteristics of extra- and intracellular responses of neurons in the AI region were studied in experiments with unanesthetized cats. It was established that auditory cortex neurons with similar best frequencies showed different forms of responses to tones of the corresponding frequency. About 40% of the auditory cortex neurons generated on responses to tone presentation. On — off and off responses were found in 27% of the neurons. Cortical neurons (27%) in which stimulation or inhibition of impulse discharge persisted throughout tone action were assigned to the tonic type group of cells. Approximately 6% of neurons in the AI region did not respond to a tone. During intracellular recording about 85% of the neurons responded to the turning on and/or off of a tone by generating an action potential followed by an IPSI. In 96% of the cortical neurons studied the IPSPs were a constant component of the intracellular responses to a tone. It is concluded that the inhibition of the impulse activity of the given neurons is of primarily a postsynaptic origin. Neurons showing one or another form of response differ from one another in the relative intensity and time characteristics of excitatory and inhibitory processes interacting on their postsynaptic membranes. In neurons of the phasic type inhibitory processes are dominant over excitatory, while excitatory processes are predominant in neurons of the tonic type.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 4, pp. 500–508, July–August, 1985.     

Responses of motor cortex neurons of the cat to auditory stimulation and their role in the instrumental food reflex

The responses of motor cortex neurons in the cat to the presentation of a single auditory click and a series of 10 clicks presented with 1,000/sec frequency were studied under conditions of chronic experiments before and after the development of an instrumental food reflex. After reflex development a single presentation of a positive conditioned stimulus (single click) markedly influenced for 7 sec the appearance of instrumental movements. At the same time, the immediate responses of motor cortex neurons to presentation of the conditioned auditory stimulus had no impact on the appearance in the motor cortex of discharges leading to the realization of instrumental movements. Consequently, motor cortex neurons do not require activation from afferent sensory inputs for the generation of such discharges. The immediate neuronal responses to conditioned stimulation did not inhibit the realization of the instrumental reflex. It is proposed that they are associated with the realization of motor function in the unconditioned defensive response evoked by the presentation of an auditory stimulus. The presence or absence of responses to auditory conditioned stimulation was dependent upon the signal meaning of the stimulus, its physical parameters, and the degree of excitability of the animal.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 4, pp. 539–550, July–August, 1985.     

Responses of neurons in the primary auditory cortex of the cat to the auditory motion stimuli with variable interaural delay

Unit responses in the primary auditory cortex of anesthetized cats to stationary and apparently moving stimuli resulted from a static and dynamically varying interaural delay (ITD) were recorded. The static stimuli consisted of binaurally presented tones and clicks. The dynamic stimuli were produced by in-phase and out-of-phase binaurally presented click trains with time-varying ITD. Sensitivity to ITDs was mostly seen in responses of the neurons with low characteristic frequency (below 2.8 kHz). All cells sampled with static stimuli responded to simulated motion. A motion effect could take the form of a difference in response magnitude depending on the direction of stimulus motion and a shift in the ITD-function opposite the direction of motion. The magnitude of motion effects was influenced by the position of motion trajectory relative to the ITD-function. The greatest motion effect was produced by motion crossing the ITD-function slopes.     

Responses of primary somatosensory cortical neurons to paired stimulation of the infraorbital nerve and vibrissae in cats

Extra- and intracellular responses of 128 neurons to paired stimulation of the infraorbital nerve and vibrissae, recorded in the projection zone of the vibrissae in cortical area SI, were studied in adult cats immobilized with tubocurarine. Conditioning stimulation completely suppressed the ability of different neurons to respond for periods of between 10 and 120 msec. The duration of the period of total suppression of test responses was shown to depend on the location of the stimulated vibrissa in the peripheral receptive field of the neurons studied. Excitatory and inhibitory responses of maximal intensity arose in the neurons to stimulation of receptive field centers. The functional role of the decrease in intensity of excitatory responses during stimulation of vibrissae located at different distances from centers of the receptive fields of cortical neurons is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 13, No. 2, pp. 117–124, April, 1981.     

Responses of neurons in cat primary somatosensory cortex to repetitive stimulation of vibrissae

Extra- and intracellular responses of neurons in the primary somatosensory cortex to repetitive mechanical stimulation of the vibrissae at different frequencies were studied in unanesthetized curarized adult cats. Unlike responses to electrical stimulation of the combined afferent input (the infraorbital nerve) spike discharges of neurons in response to vibrissal stimulation can reproduce rather higher frequencies of stimulation and their initial character changes more often in the course of the repetitive series. Most cortical neurons were characterized by limitation of the area of their peripheral receptive fields with an increase in the frequency of adequate repetitive stimulation. A group of cortical neurons was distinguished by its ability to respond to high-frequency stimulation and to generate burst discharges. Comparison of the frequency characteristics of spike responses of these cells and of inhibitory synaptic action in other cortical neurons led to the conclusion that this group of cells thus distinguished may be inhibitory cortical neurons. The role of interaction between excitatory and inhibitory processes arising in cortical neurons during repetitive stimulation of different areas of their receptive fields is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 14, No. 2, pp. 164–171, March–April, 1982.     

Response of parietal association cortex neurons to acoustic stimulation before and after auditory cortex blockage in the cat

The effect of auditory cortex blockade on response patterns of parietal association cortex neurons responding to different frequency tones was investigated in the cat. Blockade was produced by two methods: bilateral isolation and application of a 6% Nembutal solution to the auditory cortex surface. Frequency threshold curves were plotted for all test neurons. The majority of test neurons (84%) displayed one or two characteristic frequencies before blockade, as against only 63% of all neurons responding following blockade. Changes also affect the range of frequencies at which the cells could respond. Virtually all test neurons responded to application of a broad spectrum of frequencies under normal conditions. After blockade of the auditory cortex 69% of neurons no longer responded to tones above 8–10 kHz. This would suggest that mainly information on high frequency tones is transmitted via the auditory cortex. The question of where acoustic information for parietal association cortex neurons mostly originates is also discussed; association thalamic nuclei are thought to be the main source.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 18, No. 3, pp. 354–360, May–June, 1986.     

Responses of caudate neurons to direct electrical stimulation of the medial geniculate body in cats

Activity of 124 neurons of the caudate nucleus during stimulation of the medial geniculate by infrequent (0.5 Hz) square electrical stimuli 0.3 msec in duration and ranging in intensity from 50 µA to 1 mA was investigated extracellularly in chronic experiments on cats. Responses were recorded from 54 neurons (43%). The main types of neuronal responses were phasic activation in the form of a single spike or spike discharge, initial activation followed by inhibition, and primary inhibition of unit activity. Responses of excitatory character predominated (81% of all responses). Their latent period varied in different neurons from 2.7 to 64 msec. Latent periods of responses of the same neuron always showed great variability, so that all the responses recorded could be considered to be orthodromic. The mode of the histogram of latent periods of excitatory responses lay between 9 and 12 msec. The latent period of the inhibitory response varied from 12 to 130 msec, and in most neurons with this type of response it was 40–60 msec. An increase in the strength of stimulation was accompanied by an increase in the regularity of the responses, an increase in the number of spikes in them, and shortening of their latent period. The character and structure of the response of the same caudate neuron to stimulation of the medial geniculate body and to presentation of clicks were usually identical. The latent period of responses to clicks was longer. The particular features of the functional connection of the medial geniculate body with the caudate nucleus as a polymodal nonspecific structure of the forebrain are discussed.     

Response of cat auditory cortex neurons to tonal stimulation during and after nembutal anesthesia

During acute experiments on 20 cats a comparative study was made of neuronal reaction to a tone, as recorded during the first few hours after administration of Nembutal and after an interval of 10–30 h. No spontaneous activity was seen in 89% of auditory cortex neurons of the anesthetized cats; these produced a sterotyped on- response to the optimal frequency tone. Late neuronal spike discharges at distinct intervals of 100–150 msec appeared in response to the setting up of acoustic stimulation after a brief latent reaction lasting 9–15 msec. It was shown that this stimulation did not produce an off-response in the cortical neurons. When the animals emerged from Nembutal anesthesia, the neurons reacted very differently to the optimal frequency tone. About 76% of the cells produced an on, on-off or off response, while about 21% responded with either tonic spike discharges or total inhibition of these throughout the acoustic stimulation. In unanesthetized cats the vast majority of AI cortical neurons were capable of reacting as long as the stimulus lasted. It is shown how this ability is lost under deep Nembutal anesthetic.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 6, pp. 728–737, November–December, 1985.     

Evoked potentials of the auditory cortex on paired stimuli

Changes in the responsiveness of the auditory cortex to an acoustic click and to direct stimulation of the medial geniculate body were studied by the method of evoked potentials in an extended experiment on cats with implanted electrodes. It is shown that the minimum interval between two stimuli for which a second click produces an EP in the auditory cortex is from 30 to 50 msec. The relative refractory period consists of two parts. The first (50–100 msec) is characterized by a rapid recovery, and the second (about 500 msec) by a slow recovery. In contrast with a click, direct stimulation of the geniculate body does not produce a refractory condition but one of facilitation. The effects of Nembutal and chloralose anesthesia and the state of alertness on the recovery of auditory cortex responsiveness were investigated. The reason for the absence and the reduction of an EP from the auditory cortex to a testing click during absolute and relative refractory periods is not a passive decrease of excitability of the usual refractory kind, but an active interplay of excitatory and inhibitory processes in the cerebral cortex, geniculate bodies, and reticular formation of the brain stem.A. A. Bogomolets' Institute of Physiology, Academy of Sciences, Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 1, No. 1, pp. 54–64, July–August, 1969.     

Responses of central auditory neurons to frequency-modulated stimuli

Unit responses of the inferior colliculi of albino rats to frequency-modulated stimuli were investigated. The number of spikes, firing pattern, and duration of the discharge were determined. Parameters of unit responses obtained with different directions of frequency modulation were compared with the results of testing the effect of constant-frequency tones on the neurons. The distinguishing features of the unit responses to stimuli with different directions of frequency modulation were compared with the characteristics of the frequency-threshold curve and the lateral inhibitory zones, taken as indices of the unit responses to constant-frequency tones. With a change in stimulus frequency upward or downward from the initial level the unit responses to both directions could be similar as regards the number of spikes per discharge and the firing pattern or could differ sharply depending on the direction or, finally, they could arise only if the stimulus frequency changed in one direction. In some cases selectivity for the direction of the change in stimulus frequency was due to the width and position of the lateral inhibitory zones. However, for one-third of the neurons tested analysis of the spatial characteristics of the excitatory and inhibitory zones alone was insufficient to explain the high selectivity in the formation of a response to a stimulus with frequency modulation in one direction only.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol.6, No.3, pp.237–245, May–June, 1974.     

Response of rabbit visual cortex neurons to direct stimulation by current of different intensities

Response was investigated in 47 rabbit visual cortex neurons to direct intracortical electrical stimulation consisting of single biphasic pulses of increasing intensities within the 150–2,700 µA range. Minimum level for generating all components of response to electrical stimulation remained similar from one neuron to the next, ranging between 150 and 600 µA. Three different patterns of dependence between numbers of spikes at different stages of response and current amplitude were identified: stable, gradual and selective patterns. Percentages of stable dependences were lower for after-discharges in comparison with initial firing (at 11 and 26% respectively) and the proportion of selective dependences were somewhat higher (at 32 and 21%). The selective dependence observed could imply the existence of optimum current levels at which peak neuronal response occurs. At the same time, duration of inhibitory pauses lengthened uniformly in the response of all test neurons during increasing intensity of the electrical stimuli. Findings obtained during this study may have a practical application in matters connected with visual prosthetics in the blind.M. V. Lomonosov State University, Moscow. Translated from Neirofiziologiya, Vol. 21, No. 2, pp. 239–247, March–April, 1989.     

Responses of cortical neurons to viscero-somatic stimulation

Responses of cortical neurons in the posterior sigmoid gyrus of cats anesthetized with chloralose to electrical stimulation of somatic and visceral nerves were recorded. Bimodal viscero-somatic neurons are predominant in this part of the cortex and some of them also respond to light. Besides the polysensory modally-specific neurons it was also possible to distinguish a group of modally nonspecific cells (27%), whose responses to different stimuli did not differ statistically from each other. Simultaneous stimulation of visceral and somatic nerves led to facilitation of activity of the long-latency neurons; this was reflected in a decrease of 10 msec in the latent period of the response and an increase in the number of spikes per discharge.A. A. Zhdanov Leningrad University. Tadjik University, Dushanbe. Translated from Neirofiziologiya, Vol. 3, No. 6, pp. 574–581, November–December, 1971.     

Responses of auditory brainstem neurons in the grassfrog to clicks

Single unit recordings were made in the dorsal medullary nucleus and in the torus semicircularis of the immobilized grassfrog. The natural calls have a periodic pulsatile structure. To investigate the coding of pulse repetition rate periodic click trains with varying pulse repetition rate and an ensemble of clicks distributed randomly in time were used as stimuli. In the dorsal medullary nucleus strong time-locking to clicks was found. Most units showed an activation followed by suppression response. Some units showed a preference for pulse repetition rates matching their low-frequency sensitivity. In the torus semicircularis part of the units showed responses similar to dorsal medullary nucleus units. Other response types were activation irrespective of pulse repetition rate, and suppression followed by activation. The responses to the two stimulus ensembles were more compatible in the dorsal medullary nucleus than in the torus semicircularis.