Connections of neurons of the cat somatosensory cortex with the thalamic relay nuclei

Of 103 neurons in the rostral part of the posterior sigmoid gyrus of the cat cortex 30 responded to stimulation of the ventro-posterolateral and ventrolateral nuclei of the thalamus (VPL and VL), 42 responded to stimulation of VL only, and 31 to stimulation of VPL only. It was shown by intracellular recording that stimulation of VPL induces a spike response with or without subsequent IPSPs in some neurons and an initial IPSP in others. The spike frequency of single neurons reached 60/sec, but the IPSP frequency never exceeded 10–20/sec. Stimulation of VL was accompanied by: a) antidromic spike responses; b) short-latency monosynaptic EPSPs and spikes capable of following a stimulation frequency of 100/sec; c) long-latency polysynaptic EPSPs and spikes appearing in response to stimulation at 4–8/sec; d) short-latency IPSPs; e) long-latency IPSPs increasing in intensity on repetition of infrequent stimuli. It is concluded that the afferent inputs from the relay nuclei to neurons of the somatosensory cortex are heterogeneous. An important role is postulated for recurrent inhibition in the genesis of the long-latency IPSPs arising in response to stimulation of VL, and for direct afferent inhibition during IPSPs evoked by stimulation of VPL. It is shown that the rostral part of the posterior sigmoid gyrus performs the role of somatic projection and motor cortex simultaneously.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 3, pp. 245–255, May–June, 1972.     

Callosal neurons: monosynaptic connection between them and their descending projections

Antidromic and monosynaptic unit responses to the stimulation of the corpus callosum and the symmetrical cortical area as well as antidromic responses to pyramidal tract and thalamic nuclei stimulation were recorded in the sensorimotor cortex of unanaesthetized rabbits. Out of 182 callosal neurones 13 exhibited transcallosal monosynaptic responses. 8 out of 56 callosal units responded antidromically to pyramidal tract or thalamic stimulation. Thus callosal neurones may be monosynaptically excited by callosal units via the corpus callosum and by the pyramidal tract units. It was also found that a pyramidal tract neurone may send a collateral through the corpus callosum and at the same time have a transcallosal monosynaptic input. The role of monosynaptic transcallosal excitation of callosal neurones is discussed.     

Elementary transcallosal connections of the rabbit sensomotor cortex]

Antidromic responses of two callosal neurones to a local electrical stimulation of the rabbit sensorimotor cortex may be recorded simultaneously with one microelectrode in the homotopic cortical area. In such recording conditions the relative amplitude of extracellularly recorded action potentials of the two neurones is determined primarily by the distance between these neurones and the electrode's tip. In response to the stimulation of the symmetrical area transcallosal monosynaptic excitation of the callosal neurone may occur; two callosal neurones may exite monosynaptically one and the same recorded neurone. The results suggest the existence of clusters or columns, formed jointly by the bodies and terminals of callosal neurones; a functional interconnection between symmetrical clusters or columns may exist, in particular a positive feedback.     

Functional maturation of the cat visual cortex during prenatal ontogenesis]

Development of the perceiving function of the visual cortex was studied on the foeti of cats in the second stage of antenatal development with intact placental blood circulation. It has been found that functional maturing of the cortical end of the visual analyser starts at the beginning of the second half of antenatal life. At this period EPs to stimulation of the optic nerve are recorded throughout the dorsal cortex of the contralateral hemisphere. At first they appear as slow three-phase (positive-negative-positive) oscillations of a small amplitude. As the foetus develops, the EP amplitude increases, and the EP configuration in the striate zone of the cortex becomes complex. Two weaks before birth, a short-latency negative wave appears against the background of the primary positive oscillation. In the last week of antenatal development of the foeti and in the first few days of the kittens life, EPs are represented in the specific zone of the visual cortex (g. lateralis) as two negative oscillations, and in the so-called associative zone (the middle part of the suprasylvian gyrus) by one long-latency high-amplitude negative oscillation which corresponds by latency to the second negative EP component in the striate cortex.     

Convergence of nociceptive information on the parabrachio-amygdala neurons in the rat

Neurones were recorded with extracellular micropipettes, in the parabrachial area located in the dorsolateral region of the pons of anaesthetized rats. All the neurones were identified by antidromic stimulation from the nucleus centralis of the amygdala. Numerous parabrachio-amygdala neurones (70%) were exclusively affected by noxious stimuli applied to several areas of the body. The rest of the neurones (30%) were not activated by any of these stimuli. The "nociceptive" neurones were classified in two groups: the neurones in the first group ("specific nociceptive", 55% of the whole population), responded to mechanical nociceptive and thermal nociceptive stimulation (threshold greater than 44 degrees C), with a strong and sustained activation. The neurones in the second group (15% of the whole population) responded by a strong inhibition to the nociceptive stimulation. Transcutaneous electrical stimulation demonstrated that the specific nociceptive parabrachial neurones received messages from A delta and C fibres. These results demonstrate that a spino-(trigemino)-ponto-amygdala nociceptive pathway exists which could be implicated in the emotional responses to noxious events.     

Endogenous dopamine modulates cortico-pallidal influence via GABA

Sprague-Dawley rats were used to study the influence of local application of antagonists of D1 and D2 receptors (SCH 23390 and raclopride, respectively) on neuronal responses in globus pallidus evoked by somatosensory cortex stimulation. SCH 23390 was found to produce a short-latency inhibition in response to cortical stimulation and to block the long-latency inhibition. Raclopride application suppressed the short-latency inhibition and revealed the long-latency inhibition in response to cortical stimulation. It is suggested that the observed phenomena are based on the modulation of GABA releasing in stria-pallidar terminals by endogenous dopamine.     

Relevance of the callosal transfer in defining the peripheral reactivity of somesthetic cortical neurones

Experiments were performed in 13 chloralose-anaesthetized, curarized cat preparations (monitoring of rectal temperature, heart rate, expired pCO2 and EEG), in order to ascertain whether, and to what extent, the reactivity to ipsilateral skin shocks of the neurones of the anterior ectosylvian and anterior suprasylvian gyri (AEG and ASG, respectively) is dependent on the callosal output of the somatosensory areas of the contralateral hemisphere. Indeed, we knew from previous experiments that a high proportion of AEG and ASG neurones having bilateral peripheral receptive fields (PRFs) can be excited by direct stimulation of the contralateral homonymous areas, and that the callosal fibres originating in the latter carry somesthetic impulses related to contralateral PRFs. A preliminary analysis was carried out on the amplitude and latency relationships between the evoked potentials (EP) recorded simultaneously from the two hemispheres and from the corpus callosum (CC) following stimulation of the forepaw of one side. The results obtained showed good correlations between the onset and development of the EPs picked up from the hemisphere ipsilateral to the stimulated skin, on the one hand, and onset and development of the EPs recorded from the contralateral hemisphere and the corpus callosum, on the other. At a further stage of the experiments, the EPs elicited upon ipsilateral and contralateral skin shocks in the AEG-ASG area have been recorded and averaged before, during and after the reversible inactivation of callosal somesthetic transmission. This was achieved by applying polarizing currents (0.2-1 mA) to the rostral portion of the CC, adequacy and reversibility of this method having been tested by observing, respectively, suppression and prompt restoration of transcallosal EPs and of the asynaptic spiking produced by cortical cells when antidromically invaded from contralateral homotopic cortex. It was seen that during CC blockade the EPs elicited in the AEG-ASG areas did not show any change either in amplitude or time-course if brought about by contralateral peripheral stimulation, whereas those evoked by ipsilateral skin shocks exhibited significant reduction, which was related to the strength of CC polarization and to the reduction of transcallosal EPs. In control experiments similar effects were observed after ablation of somatosensory areas of the hemisphere which send off somesthetic callosal impulses, whereas strychninization of these areas caused effects opposite in sign, i.e., enhancement of the ipsilateral but not of the contralateral EPs in the areas of the untreated hemisphere. By testing the effects of CC polarization on single AEG-ASG neurones, it was observed that the responses of units linked only with contralateral PRFs (Group I; 7 units tested) were unaffected by callosal polarization. The discharges of neurones provided with wide and bilateral PRFs (Group II; 27 units tested) were not affected if elicited by contralateral PRF shocks but were deeply impaired (in 11 neurones out the 27) when provoked by ipsilateral PRF stimulation. The effect consisted chiefly of the disappearance of the first high peak of the PSTHs, and when recording intracellularly graded events, it was mirrored by a large decrease of the postsynaptic excitatory potentials elicited in Group II neurones by ipsilateral PRF shocks. A late scattered histographic component was identified in the PSTHs of such cells, which did not appear to be significantly altered during CC blockade. These effects were observed on the ipsilateral responses of 11 out of the 27 Group II neurones so tested whereas the ipsilateral PSTHs of the remaining 16 Group II neurones either did not undergo significant changes during the callosal blockade or escaped evaluation because of high spontaneous shifts of neural responsiveness. The results are discussed mainly with a view to the possible functional role of the specific somesthetic callosal fibres in defining ipsilateral reactivity for the wide-field cells of the AEG-ASG area.     

Raphe Stimulation-Evoked Modulation of Postsynaptic Responses by Neurons of the Cat Somatosensory Cortex Activated by Stimulation of Nociceptors

We studied the effects of electrical stimulation of the raphe nuclei (RN) of the cat brain on postsynaptic potentials developing in somatosensory cortex neurons activated by nociceptive influences. Intracellular records were obtained from 15 cells, which were either selectively excited by stimulation of nociceptors (intense electrical stimulation of the dental pulp) or activated by both the above nociceptive and non-nociceptive (moderate stimulations of the infraorbital nerve or thalamic ventroposteromedial nucleus, VPMN) influences. In neurons of both groups, stimulation of both nociceptive afferents and the VPMN evoked complex responses (EPSP–AP–IPSP; IPSPs were 200 to 300 msec long). In some studied cortical neurons, isolated electrical stimulation of the RN (which caused the release of serotonin, 5-HT, in the cortex) resulted in relatively short-latency synaptic excitation, while inhibition was observed in other cells. In the case where stimulation of the RN was used as conditioning influence, such stimulation (independently of the kind of the initial response to RN stimulation) led to long-latency and long-lasting suppression of all components of the synaptic reactions evoked by excitation of nociceptors. The maximum of inhibition was observed at test intervals of 300 to 800 msec. The mechanisms underlying modulatory influences coming from the 5-HT-ergic brainstem system to neurons of the somatosensory cortex, which are activated by excitation of high-threshold (nociceptive) afferent inputs, are discussed.     

Single unit responses of the reticular and ventral anterior nuclei of the thalamus to electrical stimulation of the centrum medianum

In acute experiments on cats anesthetized with thiopental (30–40 mg/kg, intraperitoneally) and immobilized with D-tubocurarine (1 mg/kg) responses of 145 neurons of the reticular and 158 neurons of the ventral anterior nuclei of the thalamus to electrical stimulation of the centrum medianum were investigated. An antidromic action potential appeared after a latent period of 0.3–2.0 msec in 4.1% of cells of the reticular nucleus and 4.4% of neurons of the ventral anterior nucleus tested in response to stimulation. The conduction velocity of antidromic excitation along axons of these neurons was 1.7–7.6 m/sec. Neurons responding with an antidromic action potential to stimulation both of the centrum medianum and of other formations were discovered, electrophysiological evidence of the ramification of such an axon. Altogether 53.8% of neurons of the reticular nucleus and 46.9% of neurons of the ventral anterior nucleus responded to stimulation of the centrum medianum by orthodromic excitation. Among neurons excited orthodromically two groups of cells were distinguished: The first group generated a discharge consisting of 6–12 action potentials with a frequency of 130–640 Hz (the duration of discharge did not exceed 60 msec), whereas the second responded with a single action potential. Inhibitory responses were observed in only 0.7% of neurons of the reticular nucleus and 4.4% of the ventral anterior nucleus tested. Afferent influences from the relay nuclei of the thalamus, lateral posterior nucleus, and motor cortex were shown to converge on neurons responding to stimulation of the centrum medianum.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 1, pp. 36–45, January–February, 1980.     

Interaction between visceral,somatic, and visual excitation on single neurons of the lateral geniculate body in cats

Acute experiments on cats anesthetized with pentobarbital and immobilized with diplacin or listhenon showed that visceral and somatic excitation may either facilitate or inhibit single unit activity in the lateral geniculate body evoked by photic stimulation. The manifestations of facilitation were: a modulatory type of enhancement of responses of silent neurons and neurons with a low level of spontaneous activity; enhancement of responses accompanied by simultaneous depression of spontaneous activity — a sensory contrast effect; enhancement of long-latency responses; appearance of a short-latency discharge from cells with an inhibitory response to light; the appearance of responses to light in neurons not responding previously or stabilization of responses in neurons responding to light irregularly. The inhibitory effects were manifested as immediate inhibition of responses, usually long-latency, and the filling up of the inhibitory pauses of the response to light with spikes, leading to a decrease in the signal-noise ratio. Somatic stimulation was more effective and more frequently evoked facilitation of responses to light (in 74% of cells). Similar results were obtained by stimulation of the mesencephalic reticular formation. Visceral excitation gave rise to facilitatory and inhibitory effects to an almost equal degree. The results show that excitation arising as the result of visceral and somatic stimulation affects the conduction of visual information in the neuronal system of the lateral geniculate body.Ivano-Frankovsk Medical Institute. Institute of Experimental Medicine, Academy of Medical Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 5, No. 6, pp. 636–643, November–December, 1973.     

The topography of functional interhemispheric asymmetry in the visual cortex]

Functional interhemispheric asymmetry was investigated by evoked potentials method in experiments on ten cats under ethaminal anaesthesia at 200 points of the visual cortex during the action of binocular and monocular photic flashes of submaximal intensity. Topographic maps have been plotted of the functional interhemispheric asymmetry. In most of the animals a hemisphere dominant and non-dominant at the given moment can be singled out. Section of the callosal body leads to reduction of the functional interhemispheric asymmetry due to a decrease of the focus of maximum activity in the dominant hemisphere and its increase in the non-dominant one. A mozaic pattern of functional interhemispheric asymmetry has been demonstrated, as expressed in the existence of zones of inverse dominance along with prevailing zones of direct dominance. Section of the callosal body produced a decrease in the area of direct dominance and an increase in that of inverse dominance. Absolute interhemispheric asymmetry was most pronounced in the central part of the visual cortex (field 18 and its medial boundary) and the relative one, on the periphery of the visual area (fields 17 and 19).     

Trace reactions following combination of a sensory stimulus with polarization of the visual region of the cortex]

Spontaneous activity and responses to photic flashes and tones of 133 neurones were recorded in the visual cortex during polarization of the same area (1.5 to 10 muA, 5 to 30 min) and after it (one to 52 min). Responses of cells to two unimodal stimuli of different parameters were analysed, of which one was presented repeatedly during the polarization ("positive"), and the other one to three times ("negative"). Depending on the previous "learning", 47.4% of the units responded after the polarization to "positive" photic stimulus and 37,8%--to acoustic stimulus. The trace effects of the stimuli pairings are reproduced in polarization after-effect by the action of the sensory signal alone. The recorded differences in the nature and duration of the reproduction of trace processes formed to an adequate and inadequate stimuli, are due to the dissimilar action of polarizing currents on neurones of the cortex cross-section and to different effectivity of the visual and non-visual influences related to it.     

Electrophysiology of the Fetal Spinal Cord : II. Interaction among peripheral inputs and recurrent inhibition

Interactions of peripheral inputs to the motoneuron of the kitten fetus as young as 3 weeks prenatal were studied by reflex discharge from the ventral root as well as by recording from single motoneurons. Facilitation was found between two synergists in fetuses 1 to 2 weeks before birth. Intracellular recording showed that the facilitation could be explained by summation of excitatory postsynaptic potentials. Inhibition was found between antagonists in the fetuses 2 to 3 weeks before birth and was accompanied by inhibitory postsynaptic potentials. Recurrent inhibition was very powerful in the fetal spinal cord as shown by large motoneuron hyperpolarization by antidromic stimulation. Cells presumed to be "Renshaw cells" and which responded to both ortho- and antidromic stimulation with repetitive firing were shown in the 2 weeks prenatal fetus. These results lead to the conclusion that there is considerable effective synaptic connection of afferent collaterals already established by the later stage of intrauterine life and that this may be achieved independently of external stimuli.     

Mechanism of expiratory muscle activation during lower thoracic spinal cord stimulation.

Lower thoracic spinal cord stimulation (SCS) may be a useful method to restore an effective cough mechanism. In dogs, two groups of studies were performed to evaluate the mechanism of the expiratory muscle activation during stimulation at the T(9)-T(10) level, which results in the greatest changes in airway pressure. In one group, expiratory muscle activation was monitored by evoked muscle compound action potentials (CAPs) from the internal intercostal muscles in the 10th, 11th, and 12th interspaces and from portions of the external oblique innervated by the L(1) and L(2) motor roots. SCS, applied with single shocks, resulted in short-latency CAPs at T(10) but not at more caudal levels. SCS resulted in long-latency CAPs at each of the more caudal caudal recording sites. Bilateral dorsal column sectioning, just below the T(11) spinal cord level, did not affect the short-latency CAPs but abolished the long-latency CAPs and also resulted in a fall in airway pressure generation. In the second group, sequential spinal root sectioning was performed to assess their individual mechanical contribution to pressure generation. Section of the ventral roots from T(8) through T(10) resulted in negligible changes, whereas section of more caudal roots resulted in a progressive reduction in pressure generation. We conclude that 1) SCS at the T(9)-T(10) level results in direct activation of spinal cord roots within two to three segments of the stimulating electrode and activation of more distal roots via spinal cord pathways, and 2) pathway activation of motor roots makes a substantial contribution to pressure generation.     

Physiology of Photoreceptor Neurons in the Abdominal Nerve Cord of the Crayfish

Nerve fibers which respond to illumination of the sixth abdominal ganglion were isolated by fine dissection from connectives at different levels in the abdominal nerve cord of the crayfish. Only a single photosensitive neuron is found in each connective; its morphological position and pattern of peripheral connections are quite constant from preparation to preparation. These cells are "primary" photoreceptor elements by the following criteria: (1) production of a graded depolarization upon illumination and (2) resetting of the sensory rhythm by interpolated antidromic impulses. They are also secondary interneurons integrating mechanical stimuli which originate from appendages of the tail. Volleys in ipsilateral afferent nerves produce short-latency graded excitatory postsynaptic potentials which initiate discharge of one or two impulses; there is also a higher threshold inhibitory pathway of longer latency and duration. Contralateral afferents mediate only inhibition. Both inhibitory pathways are effective against both spontaneous and evoked discharges. In the dark, spontaneous impulses arise at frequencies between 5 and 15 per second with fairly constant intervals if afferent roots are cut. Since this discharge rhythm is reset by antidromic or orthodromic impulses, it is concluded that an endogenous pacemaker potential is involved. It is postulated that the increase in discharge frequency caused by illumination increases the probability that an inhibitory signal of peripheral origin will be detected.     

The inhibition of medullary reticulospinal neurons during the excitation of the dorsolateral portions of the pons blocking movement and muscle tonus in rats]

Hypothalamic stimulation increasing the muscle tone in hindlimbs, and excitation of the pontine dorsolateral areas inhibiting movements and the muscle tone in rats, were studied. Hypothalamic stimulation made 36.7% of the reticulospinal neurones to discharge in the form of short-latency spikes and to increase the muscle tone. The reticulospinal cells were completely inhibited by electrical stimulation of the pontine dorsolateral areas. 23.4% of the neurones only responded to the stimulation of the pontine dorsolateral areas. 35.9% of the cells did not respond to the stimulations at all. Excitation of the pontine inhibitory areas seems to prevent the descending activating effects from the brain rostral structures to spinal motor centres.     

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.     

Correlation between conduction velocities in optic nerve and optic radiation fibers in the cat

Responses of relay neurons of the dorsal lateral geniculate body to stimulation of area 17 of the visual cortex and the optic chiasma were studied in curarized cats. A high degree of correlation was found between the latent periods of antidromic responses of these neurons to stimulation of the visual cortex and orthodromic responses of the same neurons to stimulation of the optic chiasma (r=0.895; P=0.01). In 9% of cases antidromic unit responses were recorded to stimulation of the optic chiasma, evidence that the optic nerve contains centrifugal fibers. The functional role of the temporal dispersion of the afferent flow in the visual system is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 6, pp. 606–612, November–December, 1978.     

Physiological properties of sympathetic preganglionic neurones in the thoracic intermediolateral nucleus of the cat

Extracellular spikes were recorded from cell bodies of sympathetic preganglionic neurones in spinal segments T1-T3 of the cat. Each neurone was identified by its antidromic response to electrical stimulation of the sympathetic chain and was found in histological sections to lie within the intermediolateral nucleus. Physiological properties studied in detail included basal activity, spike configuration, and latency of antidromic activation. Also studied, in tests with paired stimuli, were the threshold interstimulus interval evoking two responses, as well as changes in amplitude and latency of the second spike which occurred at intervals near this threshold. Approximately 60% of the units studied were spontaneously active, the rest were silent. Spontaneous activity was characterized by a slow (mean = 3.1 +/- 2.6 (SD) spikes/s), irregular pattern of discharge. With approximately one-third of the cases there was a periodic pattern of discharge in phase with oscillations in blood pressure. This correlation of phasic activity suggests that many of the units studied were involved specifically in cardiovascular function. Silent and spontaneously active units could not be differentiated on the basis of latency of antidromic activation or threshold interstimulus interval; mean latency for the two groups was 7.2 +/- 4.9 ms, mean threshold interval was 6.4 +/- 4.7 ms. Thus, with the exception of basal activity, the physiological properties studied failed to indicate more than a single population of neurones. These results therefore suggest that the sympathetic preganglionic neurones in the intermediolateral nucleus subserving varied autonomic functions share overlapping physiological properties, and that functional differentiation of these neurones may be based on differences in synaptic inputs.     

Centrifugal and centripetal connections of the cat visual cortex

In experiments on curarized cats unit responses in the dorsal lateral geniculate body to stimulation of various zones in area 17 of the visual cortex were analyzed. Of all cells tested 69% were found to respond antidromically and 8% orthodromically; in 7.6% of cells IPSPs occurred either after an initial antidromic spike or without it. The velocities of conduction of excitation along the corticopetal fibers of the optic radiation varied from 28 to 4.3 m/sec, but the three commonest groups of fibers had conduction velocities of 28–19, 14–12, and 10–9.5 m/sec. A difference between latent periods of antidromic responses of the same neurons was found to stimulation of different zones of the visual cortex; this indicates that axons of geniculo-cortical fibers split into several branches which form contacts with several neurons in area 17 of the visual cortex. The degree and possible mechanisms of cortical influences on neurons of the lateral geniculate body are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 3, pp. 243–249, May–June, 1976.