The dynamics of the activating and inhibitory types of cortical neuron synchronization during the realization of a defensive reflex and internal inhibition

In the visual and sensorimotor areas of the neocortex and in the hippocampus of alert nonimmobilized rabbits, in response to combinations of light flashes with electrocutaneous limb stimulation an increase was observed of synchronization in the activity of the near-by neurones by activation by inhibitory type (coincidence of the presence and absence of impulse activity). In response to flashes against the light background--conditioned inhibitor--in the visual cortex synchronization of neurones increased by inhibitory type, and in the sensorimotor cortex and hippocampus changes of synchronization appeared, similar to the action of pain reinforcement but considerably weaker. The increase of synchronization by the activation type took place mainly in the neurones pairs with unidirected increase of impulses frequency and by the inhibitory one--with its decrease. Along with this, in a considerable part of neurones pairs both changes of synchronization appeared at the impulses frequency changes of different direction.     

Effect of substance P on behavior in the rabbit and the activity of cortical neurons during training

Subcutaneous injection of substance P to rabbits in a dose of 250 mcg/kg elicited a transitory disappearance of motor reactions to painful reinforcing stimuli and a reduction of their probability to reinforced and inhibitory light flashes, as well as a protracted heart rate increase and decrease of respiration rate. One third of the neurones recorded decreased their background firing level and or excitatory components of the reactions to reinforcement and conditioned light flashes. The decrease was most distinctly seen in the sensorimotor cortex and less pronounced in the visual cortical area and hippocampus. The influence of the substance P on different types of cortical inhibition was not the same. Tonic inhibition of neuronal activity in response to reinforcement was enhanced. Bioelectrical parameters which reflect an enhancement of inhibitory hyperpolarization during elaboration of internal inhibition (i.e. inhibitory firing delays and corresponding background and evoked slow potentials oscillations) were not changed.     

Cortical network activity before behavioral reaction and before its omission in rabbits with defensive dominanta

Defensive dominanta was formed in rabbit CNS. Activity of the cortical neuronal network was investigated in these rabbits in the state of quiet wakefulness and in the intervals between the presentations of testing stimulus (light flashes). Statistical analysis of spike trains revealed some distinctions in neuronal functional organizations in the excitation focus (sensorimotor cortex) and in the visual cortex in the states of quiet wakefulness, before the movement of the paw, and before the omission of the reaction. The evidence of different roles in the network activity of sensorimotor neurons that responded and not responded to light was obtained.     

Effect of the enkephalin derivative DAGO on neuronal behavior and activity in the neocortex and hippocampus of rabbits during the acquisition of conditioned defense and inhibitory reflexes

After subcutaneous injection of 25 mkg/kg morphine-like opiate--DAGO a decrease was observed of probabilities of rabbits movements at light flashes--defensive reflex signal. The level of the background neuronal impulse activity became gradually lower in the sensorimotor cortex and the hippocampus and did not change in the visual cortex. Decrease and restoration of responses to the reinforcing stimulus (electrocutaneous limb stimulation) in all studied cortical zones proceeded in one direction while there were significant differences in dynamics of responses to inhibitory and reinforced light flashes depending on the studied cortical zone and biological significance of the stimulus. Appearance is discussed of particular characteristics of neurones systemic organization during learning at change of reinforcement properties under the influence of the studied substance, as well as similarity of some features of mechanisms of internal inhibition elaboration in a defensive situation and of properties of positive reinforcement.     

Effect of ethanol on excitatory and inhibitory processes in the visual cortex of rabbits undergoing learning

Subcutaneous ethanol injection to waking rabbits in a dose of 2-6 g/kg causes discoordination of movements, a decline in motor reactions to inhibitory and reinforced light flashes, and appearance of equalizing and paradoxical relations in the number of reactions to these stimuli. Under the influence of ethanol, activating and disinhibitory action of the pain reinforcement on neurones of the visual area temporarily weakens, while the disinhibitory influence of light flashes (CS) is preserved. Judging from the dynamics of phasic reactions to the inhibitory light flashes, the ethanol in a dose of 2-6 g/kg does not influence the inhibitory hyperpolarization processes in the cerebral cortex of rabbits. Tonic inhibition of the cortical unit activity temporarily intensifies after the ethanol injection.     

Effect of piracetam on the neuronal activity of the neocortex and on behavior in rabbits during learning

Against the background of the action of piracetam--a cyclic derivative of GABA--in a dose of 200-400 mg/kg, no significant changes were observed of probabilities of motor reactions to inhibitory and reinforced light flashes. Piracetam in that dose did not affect inhibitory pauses in responses of neurones in the visual area and corresponding late components of the evoked potential to nonreinforced light flashes, i.e. it did not intensify inhibitory hyperpolarization processes in the cerebral cortex. Piracetam administration improved differentiation of inhibitory and reinforced light flashes judging by bioelectric parameters of the brain activity as a result of intensification of pain reinforcement action on cortical neurones. The carried-out experiments revealed significant differences in neurophysiological mechanisms of action of piracetam and fenibut--GABA linear derivate related to nootropic class.     

Correlation between activities of neurons of sensorimotor and visual cortices after forming a hidden focus of excitation (defensive dominanta) in cortical representation of a forelimb in rabbits

The interaction between neurons of sensorimotor and visual cortices was investigated by cross-correlation analysis. In this interaction, we examined the role of sensorimotor neurons responding to light. In rabbits with a hidden focus of excitation, neurons of the sensorimotor cortex responding to light significantly more often formed correlation joints with cells of the visual cortex than neurons not responding to light. On the other hand, neurons of the visual cortex significantly more often formed correlation joints with neurons of the sensorimotor cortex not responding to light.     

Change in reactions of visual cortex neurons to light flash upon hypothalamic and reticular stimulation

The effect of electrical stimulation of the hypothalamic positive reinforcement zone (PRZ), the neutral hypothalamic zone (NZ), and the reticular formation (RF), of the midbrain on the impulse activity of single neurons of the visual cortex evoked by light flashes was studied in unanesthetized and uncurarized white rats. Poststimulus histograms of the neurons's responses were compared. Under the influence of stimulation the evoked activity of the neurons which responded to light changed in a majority of them (from 63% for the NZ to 82% for the RF). Qualitative differences were found in the effects of PRZ and RF stimulation. The effects of PRZ stimulation consisted in the invariability or in a decrease in the frequency of the neuron's discharge comprising the short-latent response, and an increase in the frequency of the discharges in the long-latent response ("positive-motivation" type). The effects of RF stimulation were manifested in intensification of the neuronal discharges in the short-latent and a decrease in the discharges in the long-latent response ("reticular" type). The responses of a considerable number of the neurons changed to the "combined" type. The data obtained indicates the different natures and mechanisms of the ascending activating effects caused by stimulation of the PRZ and the RF and which brings about positive nonspecific reinforcements. These effects, which are manifested differently in different periods of the neuron's working cycle cannot be explained by "energizing" or "tonus" concepts; their explanation must be sought for in informational (systemic) concepts.N. I. Grashchenkov Laboratory of Problems of Functional Control in the Human and Animal Organism, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 3, No. 4, pp. 359–368, July–August, 1971.     

Responses of sensorimotor and visual cortical neurons to light in rabbits with a hidden focus of excitation in the CNS (defensive dominanta)

In the sensorimotor cortex of rabbits with a hidden focus of excitation in the CNS, the firing rate of neurons that responded to light was significantly lower (p = 0.01) than the firing rate of neurons that did not respond to light. The same phenomenon was observed in the visual cortex of intact rabbits. Both in intact rabbits and animals with the hidden focus of excitation, 36% of neurons in the sensorimotor responded to a nonspecific for them light stimulation. In the sensorimotor cortex of rabbits with the hidden focus of excitation, more (p = 0.01) neurons responded to light with the latency lower that 100 ms and less (p = 0.02) neurons responded to light with the latency from 200 to 300 ms as compared to intact animals. In the visual cortex of rabbits with the hidden excitation focus, less (p = 0.01) neurons responded to light stimulation with the latency from 50 to 100 ms as compared to intact rabbits.     

Polyfunctional character of responses of single neurons of the visual cortex of wakeful rats to light flashes

The activity of single neurons of the visual cortex in the initial state and upon presentation of a certain program of stimuli, which included a series of modality-specific (light flashes, continuous light) and nonspecific (clicks, tone) stimuli used separately and in combination, was recorded extracellularly by glass electrodes in unanesthetized and uncurarized white rats restrained in a stall. The responses of the neurons to flashes and clicks were analyzed by the poststimulus histogram method. The regular shifts of neuronal activity in response to light flashes (with a frequency of one per second) in the form of an increase or decrease of firing rate were noted not only during the first 150–200 msec (short-latent responses — SLR) but also later, after 700–800 msec (long-latent responses — LLR). The LLR differed from the SLR also by greater variability (decrease or increase upon repeating the stimuli) and by pronounced interaction with the modality-nonspecific stimuli, which had a weak effect on the SLR and by themselves very rarely evoked responses of the visual cortex neurons. The neuron could demonstrate several LLR with a different latent period. The independent nature of each LLR was indicated by the relative independence of its dynamics. All these data permit the consideration that one and the same neuron in one cycle of its activity can be included in different functional systems of the brain, which evidently provide direct reception of information arriving over specific sensory conductors and its subsequent processing. Therefore, neurons, which made up more than half of those investigated, can be regarded as polyfunctional.N. I. Grashchekov Laboratory of Problems of Controlling Functions in Man and Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 2, No. 3, pp. 242–250, May–June, 1970.     

On neurotransmitter mechanisms of reinforcement and internal inhibition

Experiments reported in this study have been performed in order to investigate cholinergic and GABA-ergic neurotransmitter systems and substance P in the realization of internal inhibition and pain reinforcement. This was accomplished during the elaboration of inhibitory and defensive conditioned reflexes to light flashes in alert, nonimmobilized rabbits. Present results together with a review of past research indicate that the cholinergic system is directly involved in transmitting the effects of pain reinforcement to neocortical neurons. Substance P, a neuropeptide, reduces the background activity of neocortical and hippocampal neurons and the response of cortical neurons to pain and positive conditioned stimuli. The cholinergic system and substance P exert a modulating effect on the elaboration of internal inhibition. Phenybut, a GABA derivative capable of penetrating the blood-brain barrier, enhances inhibitory hyperpolarization in the cerebral cortex and improves discrimination between the inhibitory and reinforcing light flashes. It appears, therefore, that the GABA-ergic system plays a leading part in the elaboration of internal inhibition. Neuronal activity and slow potential changes in response to positive conditioned and pain stimuli occur in the same direction after administering the preparations, and the dynamics of these changes is different from that in responses to inhibitory stimuli. It may be supposed on these grounds that the neurotransmitter and neuromodulator systems studied possess a considerable degree of plasticity.     

Ensemble activity of neurons in the visual, frontal, and sensorimotor cortices and dorsal striatum during actualization of behavioral program under conditions of strategy choice

Unit and network activity of neurons in the visual, sensorimotor, and frontal cortical areas and dorsal striatum was investigated in cats under conditions of choice of the reinforcement value depending on its delay. The animals did not differ from each other in behavior. After immediate or delayed responses cats got low- or highly-valuable reinforcement, respectively. Single-unit activity in the visual and sensorimotor cortical areas and dorsal striatum was similar during performance of immediate and delayed responses. However, significant inhibition was observed in the frontal neurons during the delay period. The network activity of visual and frontal cortex displayed smaller number of interneuronal interactions during delayed responses as compared to immediate reactions. The network activity of neurons in the brain structures under study pointed to the interstructural interaction, but only during delayed reactions, steady interneuronal communication was observed between the frontal cortex and dorsal striatum. Thus, both types of estimation of cellular activity revealed differences in the ensemble organization during different types of behavior and showed specific reactions of neuronal ensembles.     

Neuronal impulse trains of the visual and sensorimotor areas of the rabbit neocortex in calm wakefulness and pseudoconditioning

The conjugation of unit activity in the neocortical visual and sensorimotor areas during calm wakefulness and in intersignal intervals, in two groups of rabbits at pseudoconditioning was studied. The first group was presented in a random order with flashes and electrocutaneous stimuli, the second one--with sounds and electrocutaneous stimuli. The number of neurones pairs working in correlation during calm wakefulness is significantly less (35%) than during pseudoconditioning (49 and 50% in the first and second rabbits groups, respectively). During calm wakefulness and in both groups during pseudoconditioning, the number of pairs with delays of discharges of the visual area neurones after the sensorimotor one, and of the sensorimotor after visual up to 120 ms was equal. Comparison of the data on delayed neuronal discharges during calm wakefulness and pseudoconditioning with those obtained earlier with conditioned reflexes testifies that forestalling of visual area neuronal discharges by sensorimotor discharges is characteristic only for the activity of cortical projections of conditioned and unconditioned stimuli.     

Changes in response of neurons in visual area of cerebral cortex of rabbits to flashes of light under the influence of low-intensity physical factors of non-ionizing nature

The effect of various physical factors (SM F: 460 O; microwave EMF: 6 GHz, continuous mode, 200 microW/sm2; sound: clicks of 50 Hz, 6 db above a threshold of EEG response) on responses of neurons in visual area of cerebral cortex of rabbits to light flashes (1 Hz, 1 ms, 0.62 J) has been studied in experiments on 27 rabbits. The character of changes depended on the indicators for a background and for the response to the isolated action of light. Inhibition, rather than activation, was observed at a significantly higher initial frequency. Effect of the factors of magnetic nature was similar to the action of sound (inadequate irritant for the visual area). Inhibitory reactions were observed more frequently (significant result for the group of neurons), with their amplification at a combined action of irritants (SMF and microwave EMF; SMF and sound). The basic character of changes was limited to the drop in the pulsation frequency at the first phase of activation and to the increase in the latent periods of the first and second active phases. Other indicators for reaction to light flashes actually didn't change.     

Fast visuomotor processing of redundant targets: the role of the right temporo-parietal junction

Parallel processing of multiple sensory stimuli is critical for efficient, successful interaction with the environment. An experimental approach to studying parallel processing in sensorimotor integration is to examine reaction times to multiple copies of the same stimulus. Reaction times to bilateral copies of light flashes are faster than to single, unilateral light flashes. These faster responses may be due to 'statistical facilitation' between independent processing streams engaged by the two copies of the light flash. On some trials, however, reaction times are faster than predicted by statistical facilitation. This indicates that a neural 'coactivation' of the two processing streams must have occurred. Here we use fMRI to investigate the neural locus of this coactivation. Subjects responded manually to the detection of unilateral light flashes presented to the left or right visual hemifield, and to the detection of bilateral light flashes. We compared the bilateral trials where subjects' reaction times exceeded the limit predicted by statistical facilitation to bilateral trials that did not exceed the limit. Activity in the right temporo-parietal junction was higher in those bilateral trials that showed coactivation than in those that did not. These results suggest the neural coactivation observed in visuomotor integration occurs at a cognitive rather than sensory or motor stage of processing.     

Dynamics of driven rhythm in neuronal assemblies in sensorimotor and visual cerebral cortices in rabbit

Coincident activity of pairs of neurons in the sensorimotor and visual areas of the cerebral cortex was studied in naive, learning, and trained rabbits during the formation of a hidden excitation focus in their central nervous system (a defensive dominanta) of the rhythmic nature. In the trained rabbits (as compared to the naive animals), percent of neuronal pairs (both neighboring and distant) in whose coincident activity the rhythm of stimulation prevailed was higher. In the visual cortex, percent of such pairs was significantly higher only for the distant neurons. Analysis of interaction between neurons in the visual and sensorimotor cortices revealed increasing the number of neuronal pairs with the driven rhythm while training. Such an increase was observed when both sensorimotor and visual neurons were considered as leading.     

Synchronization dynamics in the neuronal work of the neocortex and hippocampus during learning before and after the administration of nootropics and narcotics

The basal difference in action of the studied drugs was that nootropics (phenybut in a dose of 40 mg/kg and pyracetam in a dose of 200-400 mg/kg) did not change the initial action of pain reinforcement on synchronism in responses of the cortical neurones of alert nonimmobilized rabbits by inhibitory type (coincidence of the presence and absence of impulse activity) towards its decrease, while narcotics of various types (ethanol in a dose of 4-6 mg/kg, morphine-like opiate DAGO and opioid peptide DADLE in doses of 250 mkg/kg) eliminated the action of pain reinforcement on synchronism in responses of the cortical neurones both by inhibitory and activation (time of coincidence only of the presence of impulse activity) types. These and other drugs mainly weakened the initial action of both the inhibitory and reinforced light flashes of synchronism in neurones activity both by inhibitory and activation types. There was no constant parallelism between changes of synchronization and the frequency of the cortical impulses.     

Conjugation of unit activity in the visual and sensomotor areas of the rabbit neocortex during a conditioned reflex to sound and light

Conjugation of unit activity in the visual and sensorimotor neocortical areas was studied by means of histograms of cross- and autocorrelation in rabbits with conditioned reflex to light (1st group) and sound (2nd group). Relative number of neurones pairs acting in correlation in the areas remote from each other, in intersignal intervals both before and after stimuli did not differ in the 1st and 2nd groups. At the same time delays in neuronal discharges in one area after the other were different. In the 1st group animals there was a predominance of the number of visual area neurones discharging after sensorimotor with a delay up to 125 ms, in comparison with the number of sensorimotor area neurones discharging after the visual one. In the 2nd group rabbits the number of visual area neurones with such a delay of discharges after sensorimotor was less and, on the contrary, a predominance of sensorimotor area neurones was observed discharging after the visual one. The obtained results allow to suggest that neurones of the visual and sensorimotor neocortex areas form a single functional system in cases when conditioned and unconditioned stimuli are addressed to these areas and when only one of the studied areas is the projection zone for the combined stimuli. Organization of the neurones activity in systems in these two cases is different.     

The Impact of Spatial Incongruence on an Auditory-Visual Illusion

Background

The sound-induced flash illusion is an auditory-visual illusion – when a single flash is presented along with two or more beeps, observers report seeing two or more flashes. Previous research has shown that the illusion gradually disappears as the temporal delay between auditory and visual stimuli increases, suggesting that the illusion is consistent with existing temporal rules of neural activation in the superior colliculus to multisensory stimuli. However little is known about the effect of spatial incongruence, and whether the illusion follows the corresponding spatial rule. If the illusion occurs less strongly when auditory and visual stimuli are separated, then integrative processes supporting the illusion must be strongly dependant on spatial congruence. In this case, the illusion would be consistent with both the spatial and temporal rules describing response properties of multisensory neurons in the superior colliculus.

Methodology/Principal Findings

The main aim of this study was to investigate the importance of spatial congruence in the flash-beep illusion. Selected combinations of one to four short flashes and zero to four short 3.5 KHz tones were presented. Observers were asked to count the number of flashes they saw. After replication of the basic illusion using centrally-presented stimuli, the auditory and visual components of the illusion stimuli were presented either both 10 degrees to the left or right of fixation (spatially congruent) or on opposite (spatially incongruent) sides, for a total separation of 20 degrees.

Conclusions/Significance

The sound-induced flash fission illusion was successfully replicated. However, when the sources of the auditory and visual stimuli were spatially separated, perception of the illusion was unaffected, suggesting that the “spatial rule” does not extend to describing behavioural responses in this illusion. We also find no evidence for an associated “fusion” illusion reportedly occurring when multiple flashes are accompanied by a single beep.     

Effect of strychninization of the rabbit visual cortex on evoked activity in the lateral geniculate body

Changes in the parameters of responses of "low-frequency" and "high-frequency" neurons evoked in the lateral geniculate body by flashes or electrical stimulation of the optic nerve were investigated in immobilized and anesthetized rabbits after strychninization of the visual cortex. Under these conditions mainly an increase in the mean discharge frequency in the responses and a decrease in their latent periods were found in the "low-frequency" neurons, probable evidence of dominance of corticofugal facilitation. In most "high-frequency" neurons, on the other hand, the mean discharge frequency in the responses decreased and the latent periods were increased, evidence of dominance of corticofugal inhibition. It is suggested that reciprocal corticofugal influences exist on neurons connected with central and peripheral channels of the visual projection pathway.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow, Translated from Neirofiziologiya, Vol. 8, No. 5, pp. 459–466, September–October, 1976.