The significance of the high-frequency components in the cortical potentials of dogs in a study of learning mechanisms

Electrical activity was studied of five different regions of dogs neocortex in inter-stimuli periods in the process of learning of motor habit of pressing the feeder pedal. Epidural electrodes were used. The processing was performed by means of correlation-spectral analysis in a wide band of 1-256 Hz. Values of cross-correlation coefficients, spectra of power, coherence and phase shifts were obtained. In the process of the habit consolidation the high frequencies power increased significantly (within the limits from 60 to 150-170 osc/sec), as well as the part of high coherence (over 0.75), falling on these limits, with low phase shifts. Relatively slow-wave oscillations (1-20 osc/sec) underwent changes of considerably lesser degree. A greater locality of high frequencies (in comparison with the traditional range of 1-20 Hz) was shown. The question of the nature of high frequency EEG components is discussed.     

High-frequency components of cortical potentials and their phase-coherent characteristics during the acquisition of motor-food conditioned reflexes in dogs

Phase-coherent characteristics of neocortex background electrical activity were studied in a wide frequency range (1-100 Hz) during interstimulus periods in the process of elaboration of motor alimentary conditioned reflexes in dogs. It was shown that the process of learning led to a considerable increase of the part of high (greater than 0.75) and significant (0.5-0.75) coherences, and also of small (0 +/- 15 degrees) and accounted (+/- 16 - +/- 30 degrees) phasic shifts as compared with the state of calm wakefulness. This increase lay mainly in the high-frequency range (40-100 Hz). The analysis of the interregional relations showed that they differed not only by values of phase-coherent characteristics, but also by distributions of the latter. The obtained data testify that the state of activation is manifested both in the rise of frequency of potentials oscillations and in the increase of their synchronization.     

Electrographic correlates of "inner states" caused by positive conditioned stimuli in the course of instrumental conditioning in dogs

Energy characteristics (power spectra) of short-term (less than 1 s) EEG-reactions were studied in dogs in the course of instrumental conditioning. These reactions were observed in different areas of the cortex during selective attention in response to positive conditioned stimuli. They immediately preceded strong blow with a paw on the pedal of feeding cup and taking the reward. The EEG power at these moments was 1.5-3 times higher than the baseline EEG power level in a prestimulus period. The high-frequency structure of corresponding EEG reactions comprised discrete individual spectral peaks both in traditional (1-30 Hz) and gamma (30-80 Hz) ranges and higher-frequency components (80-200 Hz) as well. In some cases, the higher-frequency components (80-200 Hz) were most pronounced.     

EEG Findings of Reduced Neural Synchronization during Visual Integration in Schizophrenia

Schizophrenia patients exhibit well-documented visual processing deficits. One area of disruption is visual integration, the ability to form global objects from local elements. However, most studies of visual integration in schizophrenia have been conducted in the context of an active attention task, which may influence the findings. In this study we examined visual integration using electroencephalography (EEG) in a passive task to elucidate neural mechanisms associated with poor visual integration. Forty-six schizophrenia patients and 30 healthy controls had EEG recorded while passively viewing figures comprised of real, illusory, or no contours. We examined visual P100, N100, and P200 event-related potential (ERP) components, as well as neural synchronization in the gamma (30-60 Hz) band assessed by the EEG phase locking factor (PLF). The N100 was significantly larger to illusory vs. no contour, and illusory vs. real contour stimuli while the P200 was larger only to real vs. illusory stimuli; there were no significant interactions with group. Compared to controls, patients failed to show increased phase locking to illusory versus no contours between 40-60 Hz. Also, controls, but not patients, had larger PLF between 30-40 Hz when viewing real vs. illusory contours. Finally, the positive symptom factor of the BPRS was negatively correlated with PLF values between 40-60 Hz to illusory stimuli, and with PLF between 30-40 Hz to real contour stimuli. These results suggest that the pattern of results across visual processing conditions is similar in patients and controls. However, patients have deficits in neural synchronization in the gamma range during basic processing of illusory contours when attentional demand is limited.     

Conditioned reactions of hippocampal neurons during amphetamine poisoning and its counteraction with haloperidol

Activity of 144 neurones of the dorsal part of the rabbits hippocamp was recorded during elaboration of motor conditioned reflex to time. Chronic amphetamine intoxication lowered the ability of hippocampal neurones to form conditioned reactions in response to pairings of sound stimuli with electrocutaneous reinforcement and fully suppressed mechanisms of reproduction by cells of engrams of previous pairings in series of their omissions Single administration of haloperidol to intact animals somewhat increased the number of neurones reacting to the pairing and their omissions in conditioned reflex to time without significantly influencing the intensity and dynamics of reproduction of endogenous cellular reactions in the series of consecutive omissions of pairing. Haloperidol administration during amphetamine intoxication elicited shifts towards normalization of conditioned activity of neurones, eliminating the suppressing action of amphetamine on mechanisms of reproduction of engrams of combined stimuli. Such "therapeutic" effect of haloperidol in many cases did not depend on the character of its psychotropic action. The properties of amphetamine and haloperidol action on the cells of the hippocamp are discussed as compared to their action on the neurones of other brain structures, previously studied in an analogous experimental situation.     

A computer analysis of EEG intersignal reactions during the the acquisition of conditioned motor-food reflexes in dogs]

Background electric activity was studied of different neocortex areas in interstimulus intervals at the stage of generalization during elaboration of motor alimentary conditioned reflexes in dogs. For this stage the appearance is typical of short-term (0.1-0.3 s) packs of high frequencies (above 40 osc./s) significantly exceeding the adjacent initial background by frequency and amplitude (along side with motor interstimulus reactions). The index of specific variation, elaborated by us, allowed to single out this EEG pattern in the initial realizations of the background activity during their input in the computer. With the purpose of further evaluation of the above phenomenon parameters non-standard approaches of computer analyses were used, directed to decomposition of the EEG curve into the system of oscillations and receipt of corresponding amplitudes frequencies distributions (maps). It was shown, the described high frequencies packs were localized on these maps in definite sufficiently compact places.     

The structure of the background EEG of the rabbit in the high-frequency portion of the spectrum]

With the purpose of studying the character and structure of high frequency bioelectric activity of rabbits cerebral cortex in the state of calm alertness, the EEG ensembles of different areas of the cortex (sensorimotor, visual, acoustic) and dorsal hippocampus were studied with FFT method. A supposition was made about the presence of systemic organization of the background EEG in rabbits cerebral cortex, reflected, in particular, in the presence of determined components both of chaotic and rhythmic character having different degrees of manifestation. Heterogeneity was revealed in distribution of energies of spectral EEG components in the studied frequency ranges from 14.7 to 100 Hz with predominance of total specific energy value in the band of 14.7-60 Hz. In coherence functions of all the studied pairs of EEG leads rhythmic component, stable in time, was absent. Functions of the mean EEG coherence in the band of 61-100 Hz had significantly greater values in comparison with the values in the band of 14.7-40 Hz.     

Correlations among cortical potentials in the dog during the elaboration of motor alimentary conditioned reflexes

Background electrocortical activity in a frequency band of 1-100 Hz of dogs with electrodes implanted in different neocortical areas was studied before and after elaboration of motor (lever pressing) alimentary conditioned responses. Conditioning led to a significant change of the distribution histograms of cross-correlation coefficients (CC) between the cortical potentials, i. e. the dispersion of CC values decreased and the level of correlation changed as well. In more tranquil animals (with lower CC values before conditioning) the level of correlation significantly increased, in excitable ones (with higher CC values before conditioning) it decreased. CC distributions after the conditioned response elaboration became similar in both cases apparently indicating the formation of a uniform state characteristic for the type of a conditioned response under study.     

Cues paired with either rapid or slower self-administered cocaine injections acquire similar conditioned rewarding properties

The faster drugs of abuse reach the brain, the more addictive they can be. It is not known why this is. Environmental stimuli associated with drugs can promote the development and persistence of addiction by invigorating and precipitating drug-seeking behaviour. We determined, therefore, whether cues associated with the self-administration of rapidly delivered cocaine (injected intravenously over 5 versus 90 seconds) would acquire greater conditioned rewarding properties, as assessed by the performance of an operant response reinforced solely by the cues. Rats nose-poked for intravenous cocaine infusions delivered either over 5 or 90 seconds. Discrete visual cues accompanied each infusion. The rats could then press a lever to obtain the cues--now a conditioned reward--or an inactive lever. Rats in both the 5- and 90-second groups pressed more on the active versus inactive lever following extensive (24 sessions) but not following limited (3 sessions) self-administration training. There were no group differences in this behaviour. Following withdrawal from cocaine self-administration, lever discrimination progressively abated in both groups and was lost by withdrawal day 30. However, the rewarding properties of the cues were not "forgotten" because on withdrawal days 32-33, amphetamine selectively enhanced active-lever pressing, and did so to a similar extent in both groups. Thus, cues paired with rapid or slower cocaine delivery acquire similar conditioned rewarding properties. We conclude, therefore, that the rapid delivery of cocaine to the brain promotes addiction by mechanisms that might not involve a greater ability of drug cues to control behaviour.     

Selective attention in dogs from energy characteristics of neocortical potentials in the 1-220 Hz band

The state of selective attention was studied in dogs in the course of instrumental conditioning. During interstimuli intervals, this state was manifested in the state of strained waiting for conditioned stimuli. Electrical activity of different areas in both hemispheres was analyzed using the Fast Fourier Transformation. It was shown that in the process of development of selective attention, the high-frequency EEG components (40-200 Hz) in the motor area of the right hemisphere and the visual and parietal areas of the left hemisphere had a predominant significance over the traditional EEG frequencies of 1-30 Hz. The state of selective attention was characterized by another functional mosaic organization of the neocortical potentials.     

Correlation of neuronal reactions of different regions of the neocortex of the rabbit to conditioned positive and inhibitory stimuli

The activity of neurones of the visual and sensorimotor areas of the neocortex was simultaneously recorded in rabbits under the action of conditioned, inhibitory stimuli (CS, IS) and at simultaneous presentation of CS and the unconditioned stimulus (US) after trans-switching of the positive and inhibitory conditioned reflexes. Mean time of conjugated reactions of simultaneously recorded pairs of neurons is similar under the action of CS, combined action of CS and US and under IS. During the IS action as compared with CS, the phasic activity of some neurones in different areas increased, and due to this the similarity of reactions also increased; in other pairs of neurones the reactions weakened and the responses similarity decreased. The frequency of the appearance of intervals of the conjugated excitation after CS corresponds to 4.3 Hz; under combined action of CS and US it increases up to 5.9 Hz, and after IS--it decreases to 3.6 Hz. The order of coming into conjugated action of different pairs of neurones changes depending on the signal significance of the stimuli.     

Regional features of electrical reactions (in the frequency band of 1-225 Hz) in the cerebral cortex to conditioned stimuli in the course of instrumental conditioning

Power spectra of short-term (less than 1 s) EEG-reactions (in the frequency band of 1-225 Hz) were studied in dogs in the course of instrumental food conditioning. These reactions were observed in different areas of the cortex in response to positive and differentiated conditioned stimuli. Regional features between the spectra were found both in the power level and frequency structure. The power of the reactions in the visual and parietal areas of the left hemisphere was higher than in the motor areas. Power spectra of reactions to differentiated stimuli were significantly lower than the spectra of reactions to positive stimuli mainly owing to the high-frequency components (80-225 Hz). In these both cases, prestimulus power spectra did not differ. The frequency structure of corresponding EEG-reactions consisted of individual spectral peaks, mainly both gamma (30-80 Hz) and higher-frequency (80-225 Hz) bands.     

Changes in the phasic activity of neuron microsystems of the somatosensory cortex of the cat during extinction of activation reactions to unreinforced stimuli

In chronic experiments on cats, three-phasic responses of neuronal microsystems in the cortical somatic area I were studied during habituation of the EEG activation reactions. Repeated stimuli of different modalities were used: electrical pulses to the forepaw, sounds, direct stimulation of the mesencephalic RF. Simultaneously with the extinction of EEG activation reactions, the three-phasic responses of the multiunit activity (MUA) also became progressively extinct: the 1st phase of primary excitation--only a little, the 2nd phase (inhibitory)--greatly, as well as the 3rd phase--the phase of secondary excitation (if it existed at the beginning). The MUA responses to all stimuli show that these neuronal microsystems are polysensory. Relatively to the nonspecific activating RF macrosystem, the investigated neuronal microsystems are autonomous because their two functionally opposed response phases--the 1st excitatory and the 2nd inhibitory--occur against the monotonous excitatory background of the EEG activation. But in some way the neuronal microsystems are connected with the RF-system because of the parallel development of the extinction process.     

Conditioned reactions to time of hypothalamic neurons. The perifornical nucleus]

In 40% of the 52 neurones of the hypothalamic perifornical nucleus in alert rabbits conditioned trace reactions of the activational (52%) and inhibitory (48%) type were recorded in the course of elaboration of a conditioned motor reflex to time. The sign and pattern of the trace responses were determined by the nature of cell reactions to actual paired stimuli. After 50 to 70 pairings, the unit trace conditioned reaction to time persisted for a period of 10 to 15 successive omissions. Trace responses were observed most frequently in the 5th of 8th omissions. In some cases conditioned enhancement of cell activity coincided with the conditioned motor response to time. This fact together with the maximal development of a summery trace cellular response at the moment of formation of conditioned motor reactions attests the participation of neurones of the perifornical nucleus in maintaining conditioned motor activity.     

Electrical reactions in the cat brain during differentiation of photic stimuli]

Study of EEG responses to photic conditioned stimuli in cats has shown that the extent of brain activation depends not on the physical strength of the stimulus, but on its biological significance. The longest activation is recorded to a positive signal, even if it is the weakest flash of light. The intensity of EEG responses to negative stimuli is determined by the nearness of their physical parameters to those of the positive signal. Repeated presentation of a non-reinforced flash leads to a gradual extinction of EEG reactions. Brain activation in response to a reinforced positive signal persists unchanged even after a large number of its repetitions. The revealed properties of brain electrical reactions to stimuli of different significance accord with those previously recorded in response to acoustic stimuli. In both cases the data are in agreement with hypothesis of a central system of signal analysis, which performs the estimation of stimuli according to their biological quality.     

Amphetamine Administration into the Ventral Striatum Facilitates Behavioral Interaction with Unconditioned Visual Signals in Rats

Background

Administration of psychomotor stimulants like amphetamine facilitates behavior in the presence of incentive distal stimuli, which have acquired the motivational properties of primary rewards through associative learning. This facilitation appears to be mediated by the mesolimbic dopamine system, which may also be involved in facilitating behavior in the presence of distal stimuli that have not been previously paired with primary rewards. However, it is unclear whether psychomotor stimulants facilitate behavioral interaction with unconditioned distal stimuli.

Principal Findings

We found that noncontingent administration of amphetamine into subregions of the rat ventral striatum, particularly in the vicinity of the medial olfactory tubercle, facilitates lever pressing followed by visual signals that had not been paired with primary rewards. Noncontingent administration of amphetamine failed to facilitate lever pressing when it was followed by either tones or delayed presentation or absence of visual signals, suggesting that visual signals are key for enhanced behavioral interaction. Systemic administration of amphetamine markedly increased locomotor activity, but did not necessarily increase lever pressing rewarded by visual signals, suggesting that lever pressing is not a byproduct of heightened locomotor activity. Lever pressing facilitated by amphetamine was reduced by co-administration of the dopamine receptor antagonists SCH 23390 (D1 selective) or sulpiride (D2 selective).

Conclusions

Our results suggest that amphetamine administration into the ventral striatum, particularly in the vicinity of the medial olfactory tubercle, activates dopaminergic mechanisms that strongly enhance behavioral interaction with unconditioned visual stimuli.     

Increased regularity of activity of cortical neurons in learning due to disinhibitory effect of reinforcement

This paper reviews the author's studies on neurophysiologic mechanisms of conditioned reflex learning. Electroencephalograms, evoked potentials, activity of neocortical and hippocampal neurons and the rabbits' behavior in the course of elaboration of defensive and inhibitory conditioned reflexes to light flashes have been recorded. Electric shock (ECS) applied to the paw served as reinforcement. The study demonstrated three types of reinforcement effect on the activity of cortical neurons: activating, disinhibitory, and inhibitory. EEG activation due to reinforcement is accompanied by a change in phasic cortical neuronal activity from chaotic or irregular, typical of rest or inhibition, to regular tonic discharges (in neocortex and hippocampus) and group discharges in the stress rhythm, 5-7 Hz in the hippocampus. Following a number of conditioning trials, the effect of reinforcement is simulated by the effect of a conditioned stimulus. With EEG activation and increased regularity in impulses, facilitation of motor reactions is observed.     

On the reactions of the human nervous system to complex-frequency optical stimulation

A computerized system for precise stimulation and analysis of electroencephalographic (EEG) reactions to two simultaneously presented frequencies of sine-wave light (one constant, 13 Hz, and the other varying from 1 to 6 Hz and vice versa) was used to study the mechanisms of human brain reactivity to complex rhythmical stimulation. The frequencies were generated by computer and presented to the subjects by three different ways: as a result of their simple summation (additively), as a product of their multiplication (multiplicatively, amplitude modulation of constant frequency by the varying frequency), or by separate presentation to different eyes. The dynamics of electroencephalograms for different types of stimulation were compared. Under all three experimental conditions, the dynamics of EEG spectra has demonstrated the same general pattern of resonance activation, which was similar to that observed for the presented signals in the case of their amplitude modulation. Significant positive shifts in the functional state of subjects were observed as a result of stimulation. The results obtained show the leading role of the processes of amplitude modulation in the interaction of integrative, adaptive, and trace mechanisms of the brain functioning during human perception of complex rhythmical stimuli.     

The impact of experimental instruction on changes in EEG power during verbal creative thinking in men and women

Features of EEG pattern during verbal creative thinking depending on experimental instruction were studied in men and women. Spectral power density was analyzed in six frequency bands (4-30 Hz). Performance of a creative task produced an increase in the power of theta (4-6 Hz) and beta2 (20-40 Hz) components and decrease in the power of alpha (8-13 Hz) and betal (13-20 Hz). Changes in the alpha and betal bands were observed, predominantly, in the posterior areas, whereas power of the thetal and beta2 bands increased in the anterior areas. Independently of instruction, women demonstrated greater synchronization in the theta1 band than men, whereas in men the desynchronization in the alpha2 band (10-13 Hz) was more pronounced. When the subjects were instructed to create original sentences, a widespread decrease in the EEG power was observed in the band of 8-30 Hz as compared to instruction "to create sentences". Thus, the instruction-related changes in EEG power were not gender-specific. They may reflect neural activity mediating selective attention.     

On the interactivity in the human visual cortex caused by specific and nonspecific inflows

Summary An interactivity (Sato, 1969) with respect not only to the interstimulus intervenal between the conditioned and test stimuli but also with respect to the time after the test stimulus was defined as a new generalized recovery function.The contour maps of the above interactivities of the cerebral evoked potentials elicited by the paired flash stimuli and by the conditioned cutaneous and test flash stimuli, respectively, were obtained in the adult human visual cortex.It was suggested to a certain degree by the autocorrelograms of background occipital EEGs and average VEPs that the more prominent VEPs are elicited, the more predominant are the alpha waves in the occipital background EEG, and vice versa. In the contour maps of the interactivity by the paired flash stimuli, cyclic changes of facilitory mountains and occlusive valleys were regular and conspicuous in the subject who exhibited predominant alpha waves in his background EEG, whereas the cyclic changes were less regular and less marked in the subject with few alpha waves in the background EEG.The cyclic changes in the former subject appeared at an interval of about 100 msec at an interstimulus interval of around 100 msec. This acts favorably in the contribution of the cyclic change in interactivity via natural stimulations to the generation of alpha waves in spontaneous EEG. The main facilitory mountains and occlusive valleys in the maps were arranged in a line parallel to the ordinate (axis of the interstimulus interval), which suggest that the conditioned flash will affect in a specific fashion each response compoment elicited via specific afferent inflows in the VEP timelocked to the test flash stimulus. On the contrary, those in the maps obtained by the conditioned cutaneous and test flash stimuli were diagonal to the abscissa (time after the test stimulus) and ordinate, and/or the peaks of facilitory mountains and bottoms of occlusive valleys tended to form a line parallel to the abscissa, which indicates that as the cutaneous somatosensory afferent inflow converges to the visual cortex nonspecific interactivities are exhibited in each component in the VEP.