Effect of local cooling of the cortex on evoked potentials in the reticular formation in response to somatosensory stimulation

Potentials evoked in nuclei of the reticular formation by electrodermal stimulation of the limbs were investigated in acute experiments on unanesthetized, immobilized rats during cooling of the somatosensory cortex in the area of representation of one forelimb. Evoked potentials in the reticular formation were found to depend on the degree of cold inhibition of the cortical primary response to the same stimulation. The peak time of the main negative wave increased from 40–50 to 60–80 msec with a simultaneous decrease in its amplitude or its total disappearance in the case of deep cooling of the cortex. Cooling of the cortex had a similar although weaker effect on the earlier wave of the evoked potential with a peak time of 14 msec, recorded in the ventral reticular nucleus. In parallel recordings of potentials evoked by stimulation of other limbs they remained unchanged at these same points of the reticular formation or were reduced in amplitude while preserving the same temporal parameters. Cooling of the cortex thus selectively delays the development and reduces the amplitude of the response to stimulation of the limb in whose area of representation transformation of the afferent signal into a corticofugal volley is blocked. Consequently the normal development of both late and early components of the potential evoked in the reticular formation by somatic stimulation requires an additional volley, descending from the cortex, and formed as a result of transformation of the same afferent signal in the corresponding point of the somatosensory cortex.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 13, No. 1, pp. 32–38, January–February, 1981.     

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.     

Spatial principle of organization of specific afferent projections and generation of evoked potentials in the somatosensory cortex

Here we investigate the functional organization of structures involved in sensory analysis in a restricted region of a cortical projection area. We have shown that stimulation of somatosensory areas I and II (SI and SII) may block an afferent volley at the level of the thalamic relay nucleus, and that SII may be selectively blocked by stimulation of SI. Also definite somatosensory connections have been demonstrated between SII, SI, and the motor cortex. We suggest that common mechanisms underlie the generation of focal reactions in projection areas of the cortex induced by stimulation of various structures. The properties of two groups of neurones from area SII are described: those having a short latency and receiving direct projections from the thalamic relay nucleus, and those of long latent period with a well-marked convergence, and reacting to stimulation of various afferent pathways. It is suggested that each path to a local point of a cortical projection areas terminates with its relay element. The signal is then directed to a common intracortical system of neurones where signals from various sources occurs (afferent, interhemispherical, subcortico-cortical, and intracortical) converge and interact. All groups of neurones are involved in the formation of the common components of evoked potentials.Presented to the All-Union Symposium: "Electrical responses of the cerebral cortex to afferent stimuli," Kiev, October, 1969.Institute of Normal and Pathological Physiology, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 2, No. 2, pp. 155–165, March–April, 1970.     

Unit responses of the first and second somatosensory areas to stimulation of the ventroposterior thalamic nucleus

Single unit responses of the first (SI) and second (SII) somatosensory areas to stimulation of the ventroposterior thalamic nucleus (VP) were investigated in cats immobilized with D-tubocurarine. In response to VP stimulation 12.0% of reacting SI neurons and 9.5% of SII neurons generated an antidromic spike. In most antidromic responses of both SI and SII neurons the latent period did not exceed 1.0 msec. The minimal latent period of spike potentials during orthodromic excitation was 1.5 msec in SI and 1.7 msec in SII. Neurons with an orthodromic spike latency of not more than 3.0 msec were more numerous in SI than those with a latency of 3.1–4.5 msec. The ratio between the numbers of neurons of these two groups in SII was the opposite. In SII there were many more neurons with a latency of 5.6–8.0 msec than in SI. EPSPs appeared after a latent period of 1.1–9.0 msec in SI and of 1.4–6.6 msec in SII. The latent period of IPSPs was 1.5–6.8 msec in SI and 2.2–9.4 msec in SII. The relative importance of different pathways for excitatory and inhibitory influences of VP on SI and SII neurons is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 2, pp. 115–121, March–April, 1976.     

Physiological mechanisms of habituation of visual evoked potentials

Experiments on cats showed that complete operative exclusion of the reticular formation by precollicular section prevents the development of habituation of evoked potentials in the primary visual projection area and lateral geniculate body. Similar results were obtained after postcollicular section of the mesencephalon. The phenomenon of habituation of visual evoked potentials is found in posttrigeminal preparations. It is postulated that the tonic inhibitory division of the reticular formation plays an important role in the development of the habituation phenomenon.Odessa State Medical Institute. Translated from Neirofiziologiya, Vol. 4, No. 5, pp. 540–544, September–October, 1972.     

Depression of evoked potentials in rat thalamic ventro-basal complex and somatosensory cortex after reticular stimulation

Experiments on unanesthetized rats immobilized with D-tubocurarine showed that electrical stimulation (100/sec) of the central gray matter and the mesencephalic and medullary reticular formation considerably depressed potentials in the somatic thalamic relay nucleus and somatosensory cortex evoked by stimulation of the forelimb or medial lemniscus. The mean threshold values of the current used for electrical stimulation of these structures did not differ significantly and were 70 (20–100), 100 (20–120), and 120 (50–200) µA, respectively. On comparison of the amplitude-temporal characteristics of inhibition of evoked potentials during electrical stimulation of the above-mentioned structures by a current of twice the threshold strength, no significant differences were found. Immediately after the end of electrical stimulation the amplitude of the cortical evolved potential and the post-synaptic components of the thalamic evoked potential was 50–60% (P<0.01) below the control values. The duration of this depression varied from 0.5 to 1 sec. An increase in the intensity of electrical stimulation of brain-stem structures to between three and five times the threshold led to depression of the presynaptic component of the thalamic evoked potential also. Depression of the evoked potential as described above was found with various ratios between the intensities of conditioning and testing stimuli.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 8, No. 5, pp. 467–475, September–October, 1976.     

Reticular unit responses in rats during functional blocking of cortical representation of the stimulated paw

Responses of single reticular units to electrodermal stimulation were studied in unanesthetized, immobilized rats during cold blocking of the cortical representation of the stimulated limbs. Local cooling of the somatosensory cortex caused reversible and opposite changes in responses of 60 of the 86 neurons tested. In 25 cells responses only to stimulation of the limb whose sensory projection was in the cooled zone were modified. In 31 neurons changes in responses to this stimulation predominated and in 22 they were comparable with changes in responses of the same neurons to electrodermal stimulation of the other limb, whose cortical representation was intact. Cold blocking of the cortical response to presentation of one of the stimuli thus modifies the conditions for information processing in the neuron net of the reticular formation selectively for the response to presentation of that same stimulus.I. M. Sechenov Institute of Evolutionary Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 13, No. 2, pp. 179–186, March–April, 1981.     

Responses in the First or Second Somatosensory Cortical Area in Cats during Transient Inactivation of the Other Ipsilateral Area with Lidocaine Hydrochloride

Simultaneous recordings were obtained from the primary and secondary somatosensory cortical areas (SI and SII) in cats anesthetized with ketamine or pentobarbital. A total of 40 individual neurons were studied (29 in SII and 11 in SI) before, during, and following injections of microliter quantities of lidocaine hydrochloride in the other ipsilateral cortical area. Activity in the cortex injected with the local anesthetic was monitored with single-neuron, multi-neuron, or evoked potential responses to determine the time course of inactivation within 0.5-2 mm of the injection sites. Recording sites in both cortical locations were in the representations of the distal forelimb. Responses were elicited by transcutaneous electrical stimulation across the receptive fields with needle electrodes. Short-latency responses were synchronously activated, and, in those circumstances where single neurons were isolated in both areas, no overall differences in latency were noted. Anesthetization of either cortical area never blocked access of somatosensory information to the intact area, even when the injected cortex was completely silenced in the vicinity of the injection mass. In 15 SII neurons and 7 SI neurons, changes were seen in short-latency evoked responses to stimulation of their receptive fields or in background activity following local anesthesia of the other area through several cycles of injection and recovery. In 7 of these 15 SII cells, changes were noted in the timing and/or firing rates of the short-latency responses; changes were noted in the short-latency responses of 2 of these 7 SI cells while SII was silenced. In 11 SII and 6 SI cells, “background” activity that was recorded during the interstimulus intervals either increased (most cases) or decreased during local anesthesia of the other area. The results are discussed in reference to the hypothesis that primary sensory cortical areas feed information forward to secondary areas, and these feed back modulatory controls to the primary regions.     

Effect of injury to the lateral reticular nucleus and nuclei of the inferior olive on electrical responses of the cerebellar cortex evoked by vagus nerve stimulation in cats

The role of the lateral reticular nucleus and nuclei of the inferior olive in the formation of cerebellar cortical evoked potentials in response to vagus nerve stimulation was determined in experiments on 28 cats anesthetized with chloralose and pentobarbital. After electrolytic destruction of the lateral reticular nucleus, in response to vagus nerve stimulation, especially ipsilateral, lengthening of the latent period and a decrease in amplitude of evoked potentials were observed; after bilateral destruction of this nucleus, evoked potentials could be completely suppressed. It is concluded that the lateral reticular nucleus relays interoceptive impulses in the vagus nerve system on to the cerebellar cortex. Additional evidence was given by the appearance of spike responses of Purkinje cells, in the form of mainly simple discharges, to stimulation of the vagus nerve. After destruction of the nuclei of the inferior olive, the latent period and the number of components of evoked potentials in response to vagus nerve stimulation remained unchanged but their amplitude was reduced. The role of the nuclei of the inferior olive as a regulator of the intensity of the flow of interoceptive impulses to the cerebellum is discussed.N. I. Pirogov Medical Institute, Vinnitsa. Translated from Neirofiziologiya, Vol. 9, No. 3, pp. 290–299, May–June, 1977.     

Reticular structures and reticulospinal pathways participating in the initiation and inhibition of spino-bulbo-spinal activity

Reflex discharges in intercostal nerves and activity of reticulospinal fibers of the ventral and lateral funiculi, evoked by stimulation of the reticular formation and of the splanchnic and intercostal nerves were investigated in cats anesthetized with chloralose (50 mg/kg). Brain-stem neuronal structures participating in the "relaying" of spino-bulbo-spinal activity were shown to lie both in the medial zones of the medullary and pontine reticular formation and in its more lateral regions; they include reticulospinal neurons and also neurons with no projection into the spinal cord. Structures whose stimulation led to prolonged (300–800 msec) inhibition of reflex spino-bulbo-spinal activity were widely represented in the brain stem, especially in the pons. Analogous inhibition of this activity was observed during conditioning stimulation of the nerves. Reticulospinal fibers of the ventral (conduction velocity 16–120 m/sec) and lateral (17–100 m/sec) funiculi were shown to be able to participate in the conduction of spino-bulbo-spinal activity to spinal neurons. In the first case fibers with conduction velocities of 40–120 m/sec were evidently most effective. Evidence was obtained that prolonged inhibition of this activity can take place at the supraspinal level.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Institute of Normal and Pathological Physiology, Slovak Academy of Sciences, Bratislava, Czechoslovakia. Translated from Neirofiziologiya, Vol. 8, No. 4, pp. 373–383, July–August, 1976.     

Role of sensory signals in the development of cortical influences on hypothalamic structures]

Neuron discharges of the hypothalamic ventro-medial and posterior nuclei were registered in immobilized cats during electrical stimulation of the ipsi- and contralateral nerves of the brachial plexus, the 1st and the 2nd somatosensory regions (SI and SII) and the visual regions of the cortex and the reticular formation (RF) of the mesencephalon. A reversible cold block of the SI and SII failed to change the effect of the nerve stimulation. Stimulation of the postero-ventral thalamic nucleus did not change the initial activity of the hypothalamic neurons and gave rise to no neuron activity that appeared on stimulation of the SII. A conclusion was drawn that cortico-hypothalamic influences that appeared on stimulation of the nerves and the postero-ventral nucleus of the thalamus were pronounced weakly.     

Role of cholinergic mechanisms in the genesis of some cortical responses

The dynamics of evoked potentials during blocking of cholinergic cortical structures was investigated in unanesthetized cats. Application of the anticholinergic drug benactyzine inhibits the negative phases of cortical responses to stimulation of the reticular formation and non-specific thalamic nuclei and also of responses to direct cortical stimulation. Direct cortical responses (DCRs), inverted by -aminobutyric acid, are also depressed, indicating the role of cholinergic mechanisms in the genesis of these responses. During blocking of cholinergic synapses, negative phases of the primary response (PR) and response to stimulation of the specific thalamic nucleus are facilitated. A tendency is then observed toward grouping of spontaneous unit discharges and abolition of inhibition of cortical neurons produced by high-frequency stimulation of the reticular formation. One cause of the increase in amplitude of the primary response (PR) to the action of anticholinergic drugs may be widening of the zone of cortical neurons involved in the response because of abolition of the localizing effect of inhibitory neurons.Institute of Physiology, Siberian Division, Academy of Sciences of the USSR, Novosibirsk. Translated from Neirofiziologiya, Vol. 2, No. 4, pp. 406–411, July–August, 1970.     

Reticular structures in the cat brain involved in startle reflexes to somatic stimuli of various modalities

Characteristics and reticular mechanisms of two types of late reflex responses, namely startle reflexes evoked by weak (tactile) and strong (high-threshold electrical stimulation of nerves) stimulation, were studied in cats anesthetized with chloralose. The peripheral and central organization of these responses, described as "tactile" and "spino-bulbo-spinal" reflexes, were subjected to comparative analysis. Definite rostro-caudal differentiation of the reticular structures involved in the formation of reflexes of these types was discovered with the aid of destructive lesions of the brain-stem reticular formation (RF). Reflexes of the first type relay mainly at the level of the pontine RF, whereas those of the second type relay mainly at the level of the medullary RF. This result was confirmed by reversible blockade of RF by injection of small doses of sodium chloride and procaine solutions into it. The functional significance of this differentiation of RF is discussed in the context of its motor and sensory functions.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 3, pp. 380–390, May–June, 1985.     

Analysis of evoked potentials from the limbic cortex

In acute experiments on cats we investigated evoked potentials from the cingulate gyrus developed in response to stimulation of somatic and visceral nerves; also potentials from various parts of the hypothalamus, and midbrain reticular formation. We showed that the nonspecific afferent system influences electrical activity in the limbic cortex through hypothalamic pathways. We consider the limbic cortex to be part of the association area of the neocortex and that the associative responses of the cortex are more complex in nature than is usually thought to be the case, and that they are formed under the influence of impulses arriving at the cortex along many specific and nonspecific pathways. The hypothalamo—cingulate system is one of the main systems of cortico—subcortical integration. It plays an important part in regulation of autonomic, somatic, and emotional responses.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 2, No. 5, pp. 451–459, September–October, 1970.     

Retrieval of delayed evoked potentials after amygdaloid stimulation

The effect of amygdaloid stimulation on retrieval of delayed evoked potentials recorded in the cortex, mesencephalic reticular formation, lateral geniculate body, and hippocampus was investigated in unanesthetized curarized cats. Delayed evoked potentials were produced to 10–400 combinations of flashes and hypothalamic stimulation and consisted of potentials arising in response to a conditioned stimulus after a delay equal to the interval between it and the unconditioned stimulus. Amygdaloid stimulation facilitated the retrieval of these potentials if they had first been extinguished or had not appeared during initial testing.Institute of Physiology, Academy of Medical Sciences of the USSR, Siberian Branch, Novosibirsk. Translated from Neirofiziologiya, Vol. 8, No. 3, pp. 300–304, May–June, 1976.     

Monosynaptic excitation of motoneurons of the accessory nerve nucleus by reticulofugal impulses

Acute experiments on cats anesthetized with chloralose and pentobarbital showed that excitation of fast-conducting (130 m/sec) reticulospinal fibers, arising during stimulation of the ipsilateral medullary reticular gigantocellular nucleus evoked monosynaptic EPSPs in motoneurons of the accessory nerve nucleus. The EPSPs had latent periods of between 0.6 and 1.0 msec (mean 0.7 msec), they reached their maximal amplitude (4.0 mV) after 2.0–2.5 msec, and lasted about 10 msec. The EPSPs underwent only weak potentiation through the different types of stimulation of the gigantocellular nucleus and were not transformed into action potentials.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 1, pp. 62–66, January–February, 1980.     

Hypothalamic evoked potentials to vagus and sciatic nerve stimulation

Hypothalamic evoked potentials to stimulation of the cervical portion of the vagus nerve and the sciatic nerve were recorded in experiments on cats anesthetized with chloralose and immobilized with succinylcholine. When both monopolar and bipolar recording techniques were used the focus of maximal activity of both "visceral" and "somatic" evoked potentials was located in the supramammillary and posterolateral region of the hypothalamus. Responses in the tuberal and anterior hypothalamus occurred in most experiments after a longer latent period, their amplitude was lower, and they were less stable. Evoked potentials in the focus of maximal activity of the posterior hypothalamus are similar in all parameters to responses of the mesencephalic reticular formation. Evoked potentials to stimulation of the visceral nerve have a higher threshold of generation and a lower amplitude than the "somatic" responses and they are inhibited more strongly when the frequency of stimulation is increased. Evoked potentials arising in the hypothalamus in response to stimulation of the vagus and sciatic nerves are regarded as nonspecific responses of reticular type.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 5, No. 3, pp. 253–260, May–June, 1973.     

Mechanisms of interaction between the parietal association and projection areas of the cat neocortex

Changes in evoked potentials in the first visual (VI), first somatic (SI), and parietal areas of the cortex during local cooling of each area were investigated under pentobarbital anesthesia. Two types of interaction were distinguished. Type I interaction was found in all areas in the early stages of local cooling and was reflected in a similar decrease in amplitude of evoked potentials in intact parts of the cortex. In the thalamic association nuclei — the pulvinar and posterolateral nucleus — somatic evoked potentials were unchanged but visual were transformed differently from those in the cortex. Type IIinteraction was found in the later stages of cooling and only between the association area and each of the projection areas. It was reflected in a greater change in amplitude of the evoked potentials and also in their configuration. In response to somatic stimulation in the early stage of type II interaction transformation of evoked potentials in the cortex took place sooner than in the nuclei; in the later stage it took place immediately after transformation of the "subcortical" evoked potentials. In response to photic stimulation transformations of cortical evoked potentials were always preceded by the corresponding transformations in the nuclei. It is suggested that type I interaction is formed by intercortical connections and type II by direct and subcortical relay connections. Differences in the role of the association area in interaction of types I and II when activated by stimuli of different modalities are discussed.Brain Institute, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 10, No. 6, pp. 573–581, November–December, 1978.     

Reticulofugal influences of interneurons in the lateral region of gray matter of the cat spinal cord

Responses of lumbar interneurons located in the most lateral regions of Rexed's laminae IV–VII to stimulation of the medial longitudinal bundle and gigantocellular reticular nucleus of medullary pyramids, red nucleus, and peripheral nerves were investigated in cats anesthetized with pentobarbital. Stimulation of the reticulospinal fibers evoked monosynaptic excitation of many interneurons specialized for transmitting activity of the lateral descending systems, but not of peripheral afferents. Convergence of excitatory influences of all three descending systems (cortico-, rubro-, and reticulospinal) was observed on some cells of this group. In addition, monosynaptic "reticular" E PSPs appeared in interneurons transmitting activity of group Ia muscle fibers and in some interneurons of the flexor reflex afferent system. Stimulation of reticulospinal fibers evoked IPSPs in some neurons of this last group. Neurons not exposed to reticulofugal influences (both specialized neurons and interneurons of segmental reflex arcs) were located chiefly in the dorsal zones of the region studied. Recordings were also obtained from single fibers of the lateral reticulospinal tracts (conduction velocity from 26 to 81 m/sec).A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 5, pp. 525–536, September–October, 1973.     

Evoked potentials to electrical stimulation of the facial nerve in the carp tectum mesencephali

Tectal evoked potentials to stimulation of the facial nerve, containing afferent fibers of nonolfactory chemoreception, in the carp are positive evoked potentials with a latent period of 5 to 25 msec which show no phase shift as the microelectrode is advanced to a depth of 600 µ. Depending on the amplitude and latency of evoked potentials seven active zones differing in one or both parameters were distinguished in the ipsilateral tectum mesencephali. The role of impulses from the medulla in the mechanism of tectal evoked potentials to facial nerve stimulation is proved by differences in latent periods and disappearance of the tectal response (although it is preserved in the primary center) after severance of connections between the two parts of the brain. Descending influences from the tectum on the primary center were found: its extirpation disturbs evoked potential generation in several parts of the medullla, so that they either disappear completely or their parameters are modified.A. A. Zhdanov State University, Leningrad. Translated from Neirofiziologiya, Vol. 8, No. 1, pp. 39–46, January–February, 1976.