Dynamics of evoked potentials in cortical projection structures during a retarded instrumental defense reflex in dogs]

Evoked potentials (EP) in the somatosensory and auditory cortical areas were studied in four dogs against the background of a retarded defensive instrumental conditioned reflex to clicks. Four phases of the reflex were singled out by the changes in the structure of EP late components (LC) in the two projection zones. The most complex LC changes in the form of intensified negative components and the appearance of additional negative LC are recorded in response to the first click of the series and to the one followed by the conditioned motor reaction (CMR). Against the background of CMR delay, the EPs increased, while during the CMR, they decreased, owing to the diminished negative components, while the positive components were preserved and sometimes intensified. An assumption has been made that cortical projection structures of paired stimuli function in one and the same regime.     

Cortical and subcortical evoked potentials and behavioral responses after destruction of the globus pallidus in CATS

Evoked potentials (EPs) to clicks were recorded in the ectosylvian and sigmoid gyri of the cortex, in the medial geniculate body, and in the median nucleus of the thalamus by means of permanently implanted electrodes in experiments on 14 cats. A motor component of the behavioral response (conditioned food reflex) developed in response to the same stimulus (clicks). In the first group of animals the region of destruction was confined to the globus pallidus. In these animals EPs to clicks appeared during the first days after the operation in all structures, although in some tests they were reduced. Meanwhile the behavioral response to clicks was absent and showed no signs of spontaneous recovery. It could be formed (retraining) later in the experiments. In the second group of animals the globus pallidus was not completely destroyed, but neighboring areas of the entopeduncular nucleus and internal capsule were coagulated. In these cats more marked changes in behavioral and electrophysiological indices were observed. EPs to clicks disappeared in some cases for 25–30 days, subsequently returning to normal. The role of the globus pallidus as a "non-specific" polymodal structure in the analysis of sensory information is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 2, pp. 141–149, March–April, 1972.     

Dynamics of evoked potentials in the auditory zone of the dog cerebral cortex following extirpation of the medial geniculate bodies during formation of goal-directed behavior]

Integrative processes in the auditory cortical zone were studied in three dogs by EEG and EP parameters during the formation of a defensive reaction to clicks (four clicks, two per second) after the extirpation of the medial geniculate bodies. In the operated animals information is transmitted to the cortical end of the auditory analyser by compensatory paths with a larger number of relays. In the auditory cortex EPs are recorded with a longer latency (12 to 16 msec), and the duration of the negative EP component is increased (up to 40 msec). It is split mainly on the ascending front. The cortical end of the analyser participates in the formation of processes of afferent synthesis. In the active period of reflex elaboration the inflow of information increases: the EP amplitude and the duration of its main negative component become greater (up to 45-55 msec), as well as the splits on its fronts. In the course of preparing for a decision, before the achievement of the conditioned reaction, a "double EP" appears, which is due to enhanced reverberation of excitations in the compensatory paths.     

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.     

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.     

Comparison of binaural release from forward masking in animals and humans. Electrophysiologic study

The EPs of the inferior colliculus and auditory cortex in anaesthetized guinea pigs and the long latency auditory EPs in alert humans were studied. The stimuli consisted of binaurally presented pairs of clicks used as a masker, and the probe, with a variable time delay between them. The greatest relative differences between out-of-phase and in-phase probe responses were observed at the beginning of the recovery course. They averaged as 1.6, 1.5 and 1.4 for the responses of the inferior colliculus, auditory cortex and long latency potentials, resp., at the stimuli intensities of 50-65 dB SPL, and then decreased to zero during the time course of the probe response recovery. Correlation of this parameter with the stimulus intensity was positive.     

Effect of extirpation of the sensorimotor cortex on the formation of conditioned reflexes to a complex signal in the ontogeny of the cat

In kittens of the age from 39 days to 4 months the influence was studied of the sensorimotor cortex (SMC) ablation on formation and preservation of previously elaborated reflex to a simultaneous complex stimulus (light + sound) with extinction of reaction to nonreinforced components of the complex signal. It is shown that SMC ablation in the age up to 2.5 months does not affect the formation or preservation of the conditioned reflex. The SMC ablation in preliminarily trained kittens in the age of 3.5 months and older leads to disinhibition of components differentiation with preservation of the positive conditioned reflex to the complex signal. In untrained kittens of the same age the SMC ablation leads to disability of animals to inhibit the motor reaction to differentiation signal. The question is discussed of SMC significance in the ontogenesis at formation of adequate forms of behaviour requiring heterosensory interaction.     

Evoked potentials during the elaboration of a conditioned reflex to hippocampal stimulation in rats

In experiments on 9 rats, the study of evoked potentials (EPs) of the CA1 field of the dorsal hippocampus to stimulation of its symmetrical part serving as a signal of drinking conditioned reflex (CR) showed that during reflex elaboration, the amplitude of the main EP components significantly decreased; CR did not appear when the population spike (PS) was absent in the hippocampal response. PS always accompanying CR was not specific only of it, it was also recorded at other behavioural reactions. Changes of fascia dentata EPs in the process of CR elaboration to stimulation of its symmetrical part consisted in decrease of the initial negative wave and increase of the positive one. The obtained data point to a significant reconstruction of excitatory and inhibitory inputs to the hippocampus and fascia dentata under the influence of conditioned activity.     

Involvement of mesencephalic reticular formation neurons in cat conditioned reflexes

Traditional defensive and operant food reflexes were used to investigate neuronal responses of the mesencephalic reticular formation. It was found that these neurons may be divided into different groups according to function, depending on how they respond to positive conditioning stimuli. Of the two main groups of neurons with sustained tonic reactions one is activated in response to positive acoustic conditioning stimulation; it no longer reacts to the same stimulus after extinction of the reflex, while the other only becomes involved in response to positive stimulation accompanying the initiation of movement. Neurons belonging to the second group begin to respond directly to acoustic stimulation after extinction of the conditioned reflex. Neurons of the mesencephalic reticular formation can thus exercise additional tonic ascending effects both in the production and inner inhibition of the conditioned reflex. The group of neurons with a phasic reaction, i.e., a double response (a direct response to sound and another produced by movement) displayed a drop in spontaneous activity during the shaping of inhibition of differentiation and of extinction in particular. It was found that the initial changes in the spike response of reticular formation neurons during conditioning and pseudo-conditioning are similar. There are thus grounds for stating that neurons of the mesencephalic reticular formation participate in the shaping, production, and inner inhibition of traditional and operant conditioned reflexes in a differentiated capacity rather than as a population reacting identically.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 18, No. 2, pp. 161–171, March–April, 1986.     

Formation of a defensive conditioned reflex to an acoustic stimulus in rabbits after early visual deprivation

The formation of a defensive conditioned reflex to sound has been studied in rabbits raised from birth up to 30 days of life in dark. It was shown that, as compared with control animals of the same age, elaboration of reflex to sound takes place in them in shorter times periods and with less pairings. This corresponds to changes in electrographic manifestations of conditioning: increased amplitude and reduced peak latency of evoked potentials to acoustic stimuli in the auditory and sensorimotor cortical zones. The data obtained testify to enahcned functional activity of the auditory cortex, apparently due to a compensatory enhancement of impulse activity coming from the intact receptors of the auditory apparatus. It has been assumed that the observed functional changes appearing in the cortical end of the signal analyser (auditory zone); in response to sound, following visual deprivation, are a consequence of an early nature training of synaptic structures with regard to perceptionof impulses of acoustic modality.     

Reproduction of electrographic correlates of a conditioned reflex after changes in the activity of the noradrenergic and dopaminergic systems

On the model of shortly delayed defensive conditioned reflex in cats, it was shown in acute experiments that pharmacologically elicited change of NA system activity (clonidine, 0.2 and 1.5 mg/kg intravenously) and of DA system activity (apomorphine, 3 mg/kg intravenously) leads to a definite manifestation of electrographic correlates of memory trace, i.e. of conditioned evoked potential (EP) in examined brain structures, as well as of conditioned neurographic response (CNR) and conditioned skin-galvanic reaction (SGR). The increase of the NA system activity causes a rise of the number of conditioned EPs in the reticular formation, hippocampus and preoptic area along with an enhancement of CNR reproduction. The increase of the DA system activity contributes to the appearance of conditioned EPs in the hippocampus, amygdalar complex and central gray matter, together with an enhancement of the reproduction of conditioned SGR.     

Neuronal activity of the cerebral cortex in cats with conditioned inhibition

Unit activity of the somatosensory cortex was studied in chronic experiments on cats during alimentary instrumental conditioned reflex performance and elaboration of conditioned inhibition (CI). First presentations of the CI signal at the beginning of CI formation caused, due to orienting reaction to a new stimulus, a levelling of the response of the neurones to the positive conditioned signal included into the CI complex. At the stage of consolidated CI, this depression proceeded gradually during the development of conditioned inhibition caused by consecutive presentations of the nonreinforced CI combination. Two groups of neurones were involved in the process of the CI performance: one of them being the same which was activated also in response to positive stimulation during performance of the conditioned response, and the second one being areactive to the positive conditioned stimulus but reducing the frequency of the background activity.     

Investigation of pallidofugal influences on hypothalamic activity

In chronic experiments on nine cats electrical stimulation of the posterior zones of the lateral hypothalamus led to reproduction of a conditioned food-getting reflex produced previously in response to clicks. During stimulation of the globus pallidus a food-getting reflex also appeared in 3 of 10 zones studied, but it did so only irregularly. In response to combined stimulation of the hypothalamus and these three zones of the globus pallidus facilitation of this reflex was observed: its intensity was increased and its latent period shortened. Stimulation of other areas of the globus pallidus inhibited the food getting reflex of hypothalamic origin on account of the intensification of purposeless motor responses. Stimulation of the globus pallidus evoked desynchronization of spontaneous hypothalamic electrical activity. After bilateral electrolytic destruction of the globus pallidus and its principal efferent pathway (the ansa lenticularis) no conditioned food-getting reflex was reproduced during hypothalamic stimulation. After pallidectomy the amplitude of the spontaneous waves and evoked potentials was reduced by a statistically significant degree in the hypothalamic zones tested. The results are evidence of the existence of regulatory functional influences of the globus pallidus on the lateral hypothalamus.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 6, No. 6, pp. 592–601, November–December, 1974.     

Functional asymmetry of the frontal cortex and lateral hypothalamus of cats during food instrumental conditioning

The synchronism and latency of auditory evoked potentials (EP) recorded in symmetric points of the frontal cortex and lateral hypothalamus of cats were measured at different stages of instrumental food conditioning and after the urgent transition to 30% reinforcement. Correlation coefficients between EPs in the cortex and hypothalamus were high (with left-side dominance) at the beginning of the experiments, when food motivation was high, and during the whole experiments in cases of high-probability of conditioned performance. Analysis of early positive P55-80 EP component showed that at all conditioning stages the peak latency of this component was shorter in the left cortical areas than in symmetrical points, whereas in the hypothalamus the shorter latency at the left side was observed at the stage of unstable conditioned reflex, and at the stage of stable reflex the latency of the studied component was shorter at the right side. During transition to 30% reinforcement, the latency was also shorter in the right hypothalamus. It is suggested that the high left-side correlation between the hypothalamus and cortex was associated with motivational and motor component of behavior rather than reflected the emotional stress induced by transition to another stereotype of food reinforcement (30%).     

Role of the parietal associative cortex in "counting" behavior of dogs

Influence of the combined and isolated lesions of areas 5 and 7 of the parietal cortex on the counting behavior was studied in experiments with 6 dogs. Instrumental feeding reaction (lifting and placing the forepaw on the foodwell) was established. The positive conditioned stimulus was a series of 5 clicks with variable interclicks intervals and the negative (non-reinforced) conditioned stimulus was a series of 3 clicks, so that asymmetrical differentiation was elaborated. Combined bilateral lesions of areas 5 and 7 and an isolated lesion of area 5 resulted in a severe impairment of the numerical discrimination for two months, whereas the isolated lesion of area 7 did not lead to any problems in differentiation. The conclusion was made that area 5 is critical for numerical discrimination of sequential stimuli.     

Reversal of signal significance of stimuli in the avoidance reflex]

The course of reversal learning in go -- no go avoidance reflex differentiation in both cats and dogs was analysed in terms of interrelations between drive and instrumental reflex activity. When signalling properties of the conditioned stimuli were reversed, marked changes in the drive state of experimental animals occurred. The increase of the fear drive influenced both the transfer of the instrumental response to the new positive conditioned stimulus and the extinction of the response to the previously positive stimulus. The quality of the conditioned stimuli and their reflexogenic strength exerted clear effects on the fear drive and on the course of reversal learning. Prefrontal lesions affected the drive state and the conditioned reflex activity of the animals during the reversal learning.     

Neuronal responses in area 7 of the cat cortex during defensive conditioning

Responses of neurons in area 7 of the parietal association cortex during and after formation of a defensive conditioned reflex to sound were recorded in waking cats. Changes in spike responses of the neurons as a result of the onset of conditioned reflex limb movements were observed in 68% of neurons. Spike responses of neurons formed as a result of learning appeared only if conditioned-reflex limb movements appeared, and they were not observed if, for some reason or other, movements were absent after presentation of the positive conditioned stimulus or on extinction of the reflex. Responses of 46% neurons to conditioned stimulation preceded the conditioned-reflex motor responses by 50–450 msec. The remaining responding neurons were recruited into the response after the beginning of movement. Characteristic spike responses of neurons to the conditioned stimulus appeared 500–900 msec before the beginning of movement and, in the case of appearance of special, "prolonged" motor responses of limb withdrawal, evoked by subsequent reinforcing stimulation.     

Role of a species-specific acoustic environment in the development of hearing in nestlings

Main parameters were studied of the acoustic evoked potentials (EPs) from L field of the caudal neostriatum of altricial nestlings of 2-8 days to pure tones in the range of species-specific signals. It has been established that auditory EPs to the tones of different frequencies differ by the terms of appearance and the degree of maturity. At every of the revealed stages of the auditory ontogenesis, including the stage of completely formed auditory sensitivity, such parameters of auditory EPs, as latencies of different phases, amplitude-temporal pattern and the course of recovery curve are different for the tones of different frequencies. The earliest to appear are the responses to the tones 0.2-4.0 kHz, corresponding to the energy of alimentary signals. Responses to tones of higher frequencies corresponding to the spectrum of other species-specific signals appear later, but the speed of their formation is higher than in the responses to the tones of low frequencies. It is suggested that the higher speed of sensitivity maturation in the high-frequency region is determined by ecologically conditioned afferentation, the function of which is fulfilled by nestlings own vocalization.     

Transformation of conditioned responses of the tonic type

A reversal of signal meaning of stimuli signalling experimental situation was performed on dogs with conditioned defensive motor reflexes to clicks: clicks and electro-cutaneous stimulation were stopped, and the animals were started to be given food in the same experimental situation. Reversal of conditioned defensive tonic stimuli to alimentary ones led to a complex of behavioral and electrographic reactions. Isolated click tests showed that signals of experimental situation considerably affect a fully-elaborated defensive motor reflex. At the same time the preparatory role of certain situational elements may be specialized with regard to the components of the conditioned reflex, in particular to the local flexor reaction.     

The dependence of the neuronal reactions of the sensorimotor cortex to a simultaneous complex stimulus on the level of the differentiation of its components]

On alert animals the change was studied of the neuronal activity of the sensorimotor cortical area of cats brain in dependence on the level of differentiation of the components of simultaneous heteromodal complex stimulus. According to the character of this dependence and a number of other parameters two groups of neurones were singled out in the sensorimotor cortex. It was shown that parameters of reactions of all recorded neurones of the sensorimotor cortex to the positive conditioned signal were the first established after consolidation of the animal conditioned motor activity. In the course of elaboration following parameters changed: expressiveness, intensity, duration and value of latency. Reactions of neurones of both groups to the inhibitory stimuli were stabilized only after consolidation of the habit of differentiation. Responses of the first group neurones changed only by the pattern of discharge, while the responses of the second group neurones could change by expressiveness of response, its sign, duration and value of latency. Oscillations of the differentiation level after finishing of the elaboration of inhibitory conditioned reactions affected only the responses of the second group neurones to complex components.