Characteristics of responses of preoptic neurons to presentation of serial cortical stimuli

Responses of neurons of the medial (MPO) and lateral (LPO) preoptic region (RPO) and adjacent hypothalamic structures to serial stimuli (6–300/sec) of the prefrontal (area 8) and cingulate (area 24) cortex, piriform lobe (periamygdaloid cortex — RPA), and hippocampus (area CA3) were investigated in acute experiments on cats under ketamine anesthesia. Four main types of responses were found: excitatory, inhibitory, excitatory on-off effect, and inhibitory on-off effect. With the use of stimuli with increasing frequencies, the direction of the response remained constant, only its intensity changed. Neurons responding to presentation of serial stimuli were localized mainly in the central part of the MPO and basal part of the LPO, where the most pronounced foci of convergence were observed. During serial stimulation of cortical structures, inhibitory responses occurred considerably more often than excitatory (ratio 3.4:1). The presence of a gradient of inhibition was established from new to old (in a phylogenetic respect) brain formations in a number of stimulated structures. In the case of stimulating the neocortex (proreal gyrus), the predominance of inhibitory responses over excitatory was minimum (1.7:1); it increased (1.9:1) in the case of stimulating the intermediate cortex (cingulate gyrus), still more (4.5:1) under conditions of stimulating the paleocortex (periamygdaloid cortex), and in the case of stimulating the archicortex (10.2:1).A. M. Gorky Medical Institute, Ukrainian Ministry of Health, Donetsk. Translated from Neirofiziologiya, Vol. 23, No. 6, pp. 720–731, November–December, 1991.     

A major role for intracortical circuits in the strength and tuning of odor-evoked excitation in olfactory cortex

In primary sensory cortices, there are two main sources of excitation: afferent sensory input relayed from the periphery and recurrent intracortical input. Untangling the functional roles of these two excitatory pathways is fundamental for understanding how cortical neurons process sensory stimuli. Odor representations in the primary olfactory (piriform) cortex depend on excitatory sensory afferents from the olfactory bulb. However, piriform cortex pyramidal cells also receive dense intracortical excitatory connections, and the relative contribution of these two pathways to odor responses is unclear. Using a combination of in vivo whole-cell voltage-clamp recording and selective synaptic silencing, we show that the recruitment of intracortical input, rather than olfactory bulb input, largely determines the strength of odor-evoked excitatory synaptic transmission in rat piriform cortical neurons. Furthermore, we find that intracortical synapses dominate odor-evoked excitatory transmission in broadly tuned neurons, whereas bulbar synapses dominate excitatory synaptic responses in more narrowly tuned neurons.     

Possible mechanisms of the anterior limbic cortex influence on the neuron activity of the vago-solitary complex

In ananesthetized cats, neurons of the nucleus of the tractus solitarius (NTS) and the dorsal motor nucleus of the vagus nerve (DMNV) revealed phasic excitatory responses to separate single vagal and cortical stimuli. Stimulation of the anterior limbic cortex combined with vagal stimulation resulted in inhibitory or excitatory modification of the vagal induced responses of the NTS and DMNV neurons. The data obtained suggest that complete inhibitory effects are related to general cortical mechanisms of control of the functional state of the brain stem visceral neurons. Selective inhibition of the vagal induced responses by limbic cortex stimulation is due to particular cortical mechanisms of the visceral sensory transmission control via the NTS neurons.     

Corticofugal influences on the neurons of different regions of the hypothalamus

Responses of the neurons of the lateral and ventromedial hypothalamic regions (HL andHvm, respectively), as well as of the area of the dorsal hypothalamus (aHd) and the projection region of the medial forelimb bundle (MFB), evoked by stimulation of the proreal cortex (field 8), cingular cortex (field 24), pyriform lobula (periamigdalar cortex), and hippocampus (CA3) were studied in acute experiments on cats under ketamine anesthesia. Distributions of the latent periods of the responses recorded from hypothalamic neurons at stimulation of the above cortical structures were analyzed. The responses were classified into primary excitatory and primary inhibitory. Stimulation of the proreal gyrus evoked four times more excitatory responses than inhibitory responses. With stimulation of the cingular gyrus, the ratio of excitatory/inhibitory responses was 1.5∶1. Stimulation of the pyriform cortex evoked activatory and inhibitory responses with a similar probability. With hippocampal stimulation, inhibitory responses appeared two times more frequently than excitatory reactions. The hypothalamus was found to be a zone of wide convergence: one-half of all responding neurons in theHL andHvm responded to stimulations of two or more tested cortical zones. In 26% of the cells, only excitatory convergence was observed, while in 10% only inhibitory convergence was found; 21% of the cells revealed mixed convergence.     

Unit responses in area 5b of the suprasylvian gyrus to stimulation of the ventral posterolateral thalamic nucleus

Unanesthetized cats were immobilized with D-tubocurarine. Single unit responses in area 5b of the suprasylvian gyrus to stimulation of the ventral posterolateral thalamic nucleus were recorded extracellularly. Of the total number of neurons tested, 32% were excited and 3% inhibited. In 65% of neurons the responses were mixed, most of them being predominantly excitatory. Repetitive stimulation of the ventral posterolateral nucleus (6–9/sec) frequently intensified the excitatory component of the responses. Sometimes inhibition, present in the response to a single stimulus, was replaced by increased excitation. However, the same response as to a single stimulus frequently appeared in response to each consecutive stimulus of a series. Stimulation of the ventral posterolateral nucleus had a mainly excitatory effect on neurons in area 5b. Stimulation of the dorsal lateral nucleus, on the other hand, inhibited their activity. This antagonism could also be observed on the same neuron. It was concluded from the short latent periods of the orthodromic responses (3–6 msec) and from the antidromic responses of the cortical neurons to stimulation of the ventral posterolateral nucleus that this nucleus has direct two-way connections with the cortex of area 5b.     

Evoked potentials and unit activity in the cat somatosensory cortex

Investigation of unit activity of the cat somatosensory cortex has shown that the principal role in the genesis of the primary response, the response to stimulation of the thalamic relay nucleus, the callosal response, and certain other forms of evoked potentials (EPs) of the somatosensory cortex is played by neurons not usually responding by spike generation during EP development. The EPs reflect what the cortical neurons received from the afferent volley, and the level of their polarization, but they are not a reliable indicator of fast nervous processes in the cerebral cortex. The EPs reflect postsynaptic potentials (PSPs) of neurons not directly participating actively in the analysis of information reaching the cortex.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No, 4, pp. 360–367, July–August, 1970.     

Comparative study of effect of cortical nucleus of amygdala and pyriform cortex on activity of bulbar respiratory neurons in cats

Comparative microelectrophysiological study of character and peculiarities of effects of the cortical nucleus of amygdala and of the periamygdalar area of pyriform cortex on impulse activity was performed on the same single functionally identified respiratory medullar neurons. A high reactivity of bulbar respiratory neurons to stimulation is established in both studied limbic structures. There is established the qualitatively different character of their response reactions at stimulation of the cortical amygdala nucleus and the periamygdalar cortex. The cortical amygdala nucleus has been shown to produce both facilitating and inhibitory effects (with predominance of the activating one) on activity of medullar respiratory neurons (without topographical orderliness). The effect of periamygdalar cortex at stimulation of various parts was characterized by topographic differentiation. The suppressing reactions of neurons in the majority of cases were recorded at stimulation of the rostral area of periamygdalar cortex, whereas the excitatory reactions-at stimulation of its caudal part. Functional organization of respiratory control of the studied limbic system structures is discussed.     

Comparative study of effects of cortical nucleus of amygdala and pyriform cortex on activity of bulbar respiratory neurons in cats

Comparative microelectrophysiological study of character and peculiarities of effects of the cortical nucleus of amygdala and of the periamygdalar area of pyriform cortex on impulse activity was performed on the same single functionally identified respiratory medullar neurons. A high reactivity of bulbar respiratory neurons on stimulation is established in both studied limbic structures. There is established the qualitatively different character of their response reactions at stimulation of the cortical amygdala nucleus and the periamygdalar cortex. The cortical amygdala nucleus has been shown to produce on the activity of medullar respiratory neurons both facilitating and inhibitory action with predominance of the activating one (without topographical orderliness). The effect of periamygdalar cortex at stimulation of various parts was characterized by topographic differentiation. The suppressing reactions of neurons in the majority of cases were recorded at stimulation of the rostral area of periamygdalar cortex, whereas the excitatory reactions--at stimulation of its caudal part. Functional organization of respiratory control of the studied limbic system structures is discussed.     

Connections of thalamic nucleus lateralis posterior with suprasylvian cortex in cats

Cats were immobilized with D-tubocurarine. Responses of 231 neurons of the thalamic nucleus lateralis posterior to cortical stimulation in areas 5b and 21 of the suprasylvian gyrus were studied. Responses of 34 neurons were antidromic, indicating the existence of a direct projection of this nucleus to the cortical areas studied. This projection was most extensive in area 5b. The long latencies (up to 60 msec) of the antidromic responses of some neurons indicate that axons of certain neurons of thalamic nucleus lateralis posterior conduct excitation very slowly (0.3 m/sec). Orthodromic responses with latencies of 2–3 msec to cortical stimulation point to the presence of direct pathways from cortex to nucleus. The flow of afferent impulses into the nucleus from area 5b is stronger than from area 21. Convergence of impulses from these areas was observed on 44% of neurons of the nucleus. Cortical stimulation of areas 5b and 21 evoked postsynaptic inhibition in most neurons of the nucleus. It is concluded that two-way direct connections exist between nucleus lateralis posterior of the thalamus and the suprasylvian cortex.     

Unit responses of the first and second somatosensory cortical areas to peripheral,thalamic, and cortical stimulation

Unit responses of the first (SI) somatosensory area of the cortex to stimulation of the second somatosensory area (SII), the ventral posterior thalamic nucleus, and the contralateral forelimb, and also unit responses in SII evoked by stimulation of SI, the ventral posterior thalamic nucleus, and the contralateral forelimb were investigated in experiments on cats immobilized with D-tubocurarine or Myo-Relaxin (succinylcholine). The results showed a substantially higher percentage of neurons in SII than in SI which responded to an afferent stimulus by excitation brought about through two or more synaptic relays in the cortex. In response to cortical stimulation antidromic and orthodromic responses appeared in SI and SII neurons, confirming the presence of two-way cortico-cortical connections. In both SI and SII intracellular recording revealed in most cases PSPs of similar character and intensity, evoked by stimulation of the cortex and nucleus in the same neuron. Latent periods of orthodromic spike responses to stimulation of nucleus and cortex in 50.5% of SI neurons and 37.1% of SII neurons differed by less than 1.0 msec. In 19.6% of SI and 41.4% of SII neurons the latent period of response to cortical stimulation was 1.6–4.7 msec shorter than the latent period of the response evoked in the same neuron by stimulation of the nucleus. It is concluded from these results that impulses from SI play an important role in the afferent activation of SII neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 4, pp. 351–357, July–August, 1976.     

The neuronal organization of the limbic (cingulo-)-visceral reflex arc

The paper summarizes new electrophysiological data concerning the structural-functional organization of the limbic cortex and role of the rostral limbic region of visceral functions. Here are presented the results of a series of electrophysiological investigation of the focus of localization in the supracallosal (area 24) and infracallosal (area 25) part of the anterior cingulate gyrus of evoked potentials of maximal amplitude and minimal latent period to stimulation of pelvic, splanchnic and sciatic nerves. It was shown that evoked potentials of maximal amplitude and minimal latent periods to stimulation of viscero-somatic nerves are recorded in the supragenual area 24 in comparison with the infragenual area 25 of the anterior limbic cortex. In a series of microelectrophysiological studies of reactions of neurons of area 24 and 25 it was established that the reactivity of neurons of area 24 is higher than that of area 25. All these data indicate to the leading role of area 24 in reception and treatment of viscero-somatic afferent signals. In series of experiments it was shown that the focus of exciting neurons, forming the descending singular-autonomic discharge is localized in the infragenual area 25 of anterior limbic cortex. In a study of the comparative characteristics of sympathetic responses in lumbar white communicating rami and parasympathetic responses in pelvic nerve it was shown that evoked potentials in pelvic nerve and white rami had the lowest threshold and shorter latency in case of stimulation of area 25. Study of characteristics of influence of dorsal (area 24) and ventral (area 25) regions of rostral limbic cortex on bioelectrical activity of two postganglionic sympathetic nerves-inferior cardiac and vertebral branches of stellate ganglion, innervating coronary vessels and vessels of anterior extremities correspondingly, showed that stimulation of ventral area 25 evoked increase of electrical activity of the two sympathetic nerves and reliable increase of systemic arterial pressure, while stimulation of dorsal area 24 evoked decrease of tonic activity of the two sympathetic nerves and reliable decrease of systemic arterial pressure. In the paper are presented also the results of microelectrophysiological investigation of peculiarities of reactions of inspiratory and expiratory neurons of bulbar respiratory center to high frequency stimulation of area 24 and 25--in case of stimulation of dorsal area 24 the prevailing effect is suppression of spike activity of neurons, of stimulation of ventral, infragenual area 25 the prevailing influence is excitatory. In another series of microelectrophysiological experiments it was shown downward blocking inhibitory influence of dorsal supragenual area 24 of anterior limbic cortex on activity of vagal viscerosensory neurons of bulbar solitary tract nucleus. It is concluded that the strictly connected one another areas 24 and 25 of limbic cortex are functionally differentiated: the infra-limbic cortex is mainly a viscero-motor cortex, while the prelimbic area 24 plays a leading role in reception and treatment of viscero-somatic afferent information.     

Cortical localization of dopamine D4 receptors in the rat brain--immunocytochemical study.

Using polyclonal antibody against dopamine D4 receptor we investigated cortical distribution of D4 receptors, with the special emphasis on regions of the prefrontal cortex. Prefrontal cortex is regarded as a target for neuroleptic drugs, and engaged in the regulation of the psychotic effects of various substances used in the experimental modeling of schizophrenia. Western blot analysis performed on samples from the rat cingulate, parietal, piriform cortices and also striatum revealed that antibody recognized one main band of approximately 40 kD, which corresponds to the predicted molecular weight of D4 receptor protein. In immunocytochemical studies we found D4 receptor-positive neurons in all regions of prefrontal cortex (cingulate, agranular/insular and orbital cortices) and all cortical regions adjacent to prefrontal cortex, such as frontal, parietal and piriform cortex. Substantial number of D4 receptor-positive neurons has also been observed within the striatum and nucleus accumbens. In general, a clear stratification of the D4 receptor-positive neurons was observed in the cortex with the highest density seen in layers II/III and V/VI. D4 immunopositive material was also found in the dendritic processes, particularly clearly visible in the layer II/III. At the cellular level D4 receptor immunoreactivity was seen predominantly on the periphery of the cell body, but a certain population of neurons with clear cytoplasmatic localization was also identified. In addition to cortical distribution of D4 receptor-positive neurons we tried also to define types of neurons expressing D4 receptor protein. In double-labeling experiments, D4 receptor protein was found in nonphosphorylated neurofilament H-positive, calbindin-D28k-positive, as well as parvalbumin-positive cells. Since, used proteins are markers of certain populations of pyramidal neurons and GABA-ergic interneurons, respectively, our data indicate that D4 receptors are located on cortical pyramidal output neurons and their dendritic processes as well as on interneurons. Above localization indicates that D4 receptors are not only directly influencing excitability of cortical inter- and output neurons but also might be engaged in dendritic spatial and temporal integration, required for the generation of axonal messages. Additionally, our data show that D4 receptors are widely distributed throughout the cortex of rat brain, and that their cortical localization exceeds the localization of dopaminergic terminals.     

Effects of Stimulation of the Periaqueductal Gray and <Emphasis Type="Italic">Locus Coeruleus</Emphasis> on Postsynaptic Reactions of Cat Somatosensory Cortex Neurons Activated by Nociceptors

In cats, we studied the influences of stimulation of the periaqueductal gray (PAG) and locus coeruleus (LC) on postsynaptic processes evoked in neurons of the somatosensory cortex by stimulation of nociceptive (intensive stimulation of the tooth pulp) and non-nociceptive (moderate stimulations of the infraorbital nerve and ventroposteromedial nucleus of the thalamus) afferent inputs. Twelve cells activated exclusively by nociceptors and 16 cells activated by both nociceptive and non-nociceptive influences (hereafter, nociceptive and convergent neurons, respectively) were recorded intracellularly. In neurons of both groups, responses to nociceptive stimulation (of sufficient intensity) looked like an EPSP-spike-IPSP (the latter, of significant duration, up to 200 msec) complex. Electrical stimulation of the PAG (which could itself evoke activation of the cortical neurons under study) resulted in long-term suppression of synaptic responses evoked by excitation of nociceptors (inhibition reached its maximum at a test interval of 600 to 800 msec). We observed a certain parallelism between conditioning influences of PAG activation and effects of systemic injections of morphine. Isolated stimulation of LC by a short high-frequency train of stimuli evoked primary excitatory responses (complex EPSPs) in a part of the examined cortical neurons, while in other cells high-amplitude and long-lasting IPSP (up to 120 msec) were observed. Independently of the type of the primary response to PAG stimulation, the latter resulted in long-term (several seconds) suppression of the responses evoked in cortical cells by stimulation of the nociceptive inputs. The mechanisms of modulatory influences coming from opioidergic and noradrenergic brain systems to somatosensory cortex neurons activated due to excitation of high-threshold (nociceptive) afferent inputs are discussed.Neirofiziologiya/Neurophysiology, Vol. 37, No. 1, pp. 61–73, January–February, 2005.     

Unit responses in area 5b of the suprasylvian gyrus to stimulation of the posterior lateral thalmamic nucleus

In response to stimulation of the posterior lateral nucleus in unanesthetized cats immobilized with D-tubocurarine an evoked potential consisting of three components with a latent period of 3–5 msec appeared in area 5b of the suprasylvian gyrus. All three components were reversed at about the same depth in the cortex (1500–1600 µ). Reversal of the potential shows that it is generated in that area by neurons evidently located in deeper layers of the cortex and is not conducted to it physically from other regions. Responses of 53 spontaneously active neurons in the same area of the cortex to stimulation of the posterior lateral nucleus were investigated. A characteristic feature of these reponses was that inhibition occurred nearly all of them. In 22 neurons the responses began with inhibition, which lasted from 30 to 400 msec. In 30 neurons inhibition appeared immediately after excitation while one neuron responded by excitation alone. The latent periods of the excitatory responses varied from 3 to 28 msec. The short latent period of the evoked potentials and of some single units responses (3–6 msec) confirms morphological evidence of direct connections between the posterior lateral nucleus and area 5b of the suprasylvian gyrus. Repetitive stimulation of that nucleus led to strengthening of both excitation and inhibition. Influences of the posterior lateral nucleus were opposite to those of the specific nuclei: the posterior ventrolateral nucleus and the lateral and medial geniculate bodies. Stimulation of the nonspecific reticular nucleus, however, evoked discharges from neurons like those produced by stimulation of the posterior lateral nucleus.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 5, pp. 502–509, September–October, 1973.     

The modulation of sensory transmission through the medullary dorsal horn during cortically driven mastication

During mastication, reflexes are modulated and sensory transmission is altered in interneurons and ascending pathways of the rostral trigeminal sensory complex. The current experiment examines the modulation of sensory transmission through the most caudal part of the trigeminal sensory system, the medullary dorsal horn, during fictive mastication produced by cortical stimulation. Extracellular single unit activity was recorded from the medullary dorsal horn, and multiple unit activity was recorded from the trigeminal motor nucleus in anesthetized, paralyzed rabbits. The masticatory area of sensorimotor cortex was stimulated to produce rhythmic activity in the trigeminal motor nucleus (fictive mastication). Activity in the dorsal horn was compared in the presence and absence of cortical stimulation. Fifty-two percent of neurons classified as low threshold and 83% of neurons receiving noxious inputs were influenced by cortical stimulation. The cortical effects were mainly inhibitory, but 21% of wide dynamic range and 6% of low threshold cells were excited by cortical stimulation. The modulation produced by cortical stimulation, whether inhibitory or excitatory, was not phasically related to the masticatory cycle. It is likely that, when masticatory movements are commanded by the sensorimotor cortex, the program includes tonic changes in sensory transmission through the medullary dorsal horn.     

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.     

Localization of the CB1 cannabinoid receptor in the rat brain. An immunohistochemical study

The presence of central cannabinoid receptor (CB1), involving the N-terminal 14 amino acid peptide, was demonstrated in the rat brain by immunohistochemistry. Intensely stained neurons were observed in the principal neurons of the hippocampus, striatum, substantia nigra, cerebellar cortex, including the Purkinje cells. Moderate CB1-IR cell bodies and fibers were present in the olfactory bulb, cingulate, entorhinal and piriform cortical areas, amygdala and nucleus accumbens. The perivascular glial fibers have shown moderate to high density CB1-IR in olfactory and limbic structures. Low density was detected in the thalamus and hypothalamus and area postrema. The CB1 receptor was widely distributed in the forebrain and sparsely in the hindbrain. These new data support the view that the endogenous cannabinoids play an important role in different neuronal functions as neuromodulators or neurotransmitters.     

Corticofugal effects on the neuronal activity in the emotiogenic/motivational zones of the hypothalamus

Neuronal responses to stimulation of the proreal (field 8) and cingular (field 24) cortices, pyriform lobe (periamygdalar cortex), and hippocampus (CA3) were studied in the lateral (HL) and ventromedial (Hvm) hypothalamus, dorsal hypothalamic region (aHd), and projection region of the medial forelimb bundle (MFB); single and repeated (series of a 6–300 sec⁻¹ frequency) stimuli were used. At single stimulations, the minimum proportion of inhibitory responses with respect to excitatory effects was observed when the neocortex (the proreal gyrus) was stimulated; this proportion became successively greater at stimulations of the intermediate cortex (the cingular gyrus) and paleocortex (the pyriform cortex), while stimulation of the archicortex (the hippocampus) evoked mostly inhibitory responses. At repeated stimulation of the cortical structures, inhibitory responses prevailed in the neurons under study: their total number was nearly four times larger than that of excitatory reactions. The response patterns to single and serial stimulations of the cortical structures allowed us to demonstrate: (i) significant diversity of the influences received by hypothalamic neurons from the cortical structures and (ii) the dependence of the pattern of these influences on the phylogenetic specificity of the above structures.     

Organization of the Connections between the Parietal Associative Zone and the Primary Sensory Zones in the Cat Neocortex

In acute experiments on cats, we studied the impulse activity of 262 neurons of the parietal associative zone (PAZ, field 5). Among them, 129 cells [100 silent units and 29 units generating background activity (BA)] were identified as output neurons, while 133 cells with the BA were interneurons of the intrinsic cortical neuronal circuits. Electrical stimulation of the primary visual, auditory, or somatosensory cortices evoked no impulse responses in silent output PAZ neurons, while output neurons with the BA and interneurons (more than 65 and 80% of the cell units, respectively) generated clear responses (more frequently, phasic). Stimulation of the auditory and visual cortices exerted mostly inhibitory effects, while stimulation of the somatosensory cortex provided mostly excitatory influences. The ratios of neurons generating primary excitatory and inhibitory responses to stimulation of the visual, auditory, and somatic cortices were 0.3:1, 0.6:1, and 3.2:1, respectively. More than 95% of the field-5 neurons were influenced from the primary sensory zones via di- and/or polysynaptic pathways. Monosynaptic excitatory inputs from the visual cortex were identified for 3.8% of interneurons and 6.9% of output PAZ neurons; for the auditory cortical inputs, the respective figures were 1.7 and 3.5%. Monosynaptic connections with the somatic cortex were found only for 4% of the interneurons under study. It has been concluded that interaction of heteromodal signals coming to the PAZ via the corticopetal and associative inputs occurs on neurons of all the cortical layers.     

The late excitatory responses of the motor cortex neurons in the cat to stimulation of the pyramidal tract

Under conditions of partial suppression of GAMKA-dependent cortical inhibition in the motor cortex of anesthetized cats, a weak electrical stimulation of the pyramidal tract evoked the late slow (50-200 ms) excitatory reactions in the motor cortex neurons similar to those previously recorded under the same conditions in response to stimulation of the parietal cortex. This finding favors the proposal that the late excitatory component of the cortico-cortical response reflects the repetitive activation of cortical neurons due to excitation spread via the system of cortical recurrent excitatory collaterals.