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.     

Imaging the spatio-temporal dynamics of supragranular activity in the rat somatosensory cortex in response to stimulation of the paws

We employed voltage-sensitive dye (VSD) imaging to investigate the spatio-temporal dynamics of the responses of the supragranular somatosensory cortex to stimulation of the four paws in urethane-anesthetized rats. We obtained the following main results. (1) Stimulation of the contralateral forepaw evoked VSD responses with greater amplitude and smaller latency than stimulation of the contralateral hindpaw, and ipsilateral VSD responses had a lower amplitude and greater latency than contralateral responses. (2) While the contralateral stimulation initially activated only one focus, the ipsilateral stimulation initially activated two foci: one focus was typically medial to the focus activated by contralateral stimulation and was stereotaxically localized in the motor cortex; the other focus was typically posterior to the focus activated by contralateral stimulation and was stereotaxically localized in the somatosensory cortex. (3) Forepaw and hindpaw somatosensory stimuli activated large areas of the sensorimotor cortex, well beyond the forepaw and hindpaw somatosensory areas of classical somatotopic maps, and forepaw stimuli activated larger cortical areas with greater activation velocity than hindpaw stimuli. (4) Stimulation of the forepaw and hindpaw evoked different cortical activation dynamics: forepaw responses displayed a clear medial directionality, whereas hindpaw responses were much more uniform in all directions. In conclusion, this work offers a complete spatio-temporal map of the supragranular VSD cortical activation in response to stimulation of the paws, showing important somatotopic differences between contralateral and ipsilateral maps as well as differences in the spatio-temporal activation dynamics in response to forepaw and hindpaw stimuli.     

Representation of auditory and somatosensory systems in the turtle thalamus: An electrophysiological study

During research on awake, curarized turtles it was found that auditory representation is located in the n. reuniens, which contains unimomodal (auditory) and bimodal (auditory and somatosensory) units. Optimal frequencies lay within the 200–400 Hz range. The somatosensory system is more extensively represented and overlaps with the auditory and lateral sections of the n. reuniens. The prevalence of contralateral somatic projections is also revealed. Test neurons could be divided between units with extensive and limited receptive fields according to the pattern of their response to presentation of auditory and somatosensory stimuli.I. S. Beritashvili Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 18, No. 4, pp. 443–453, July–August, 1986.     

Responses of units in the magnocellular part of the medial geniculate body to acoustic and somatosensory stimulation

Responses of 150 neurons in the magnocellular part of the medial geniculate body to clicks and to electrodermal stimulation of the contralateral forelimb were investigated in cats immobilized with myorelaxin. Of the total number of neurons 65% were bimodal, 16.6% responded only to clicks, and 15.4% only to electrodermal stimulation. The unitary responses were excitatory (spike potentials) and inhibitory (inhibition of spontaneous activity). Responses beginning with excitation occurred more frequently to stimulation by clicks than to electrodermal stimulation, whereas initial inhibition occurred more often to electrodermal stimulation. The latent period of the initial spike potentials in response to clicks and to electrodermal stimulation was 5–27 and 6–33 (mean 11.6 and 16.2) msec respectively. Positive correlation was found between the latent periods of spike potentials recorded in the same neurons in response to clicks and to electrodermal stimulation, and also to electrodermal stimulation and to stimulation of the dorsal funiculus of the spinal cord. It is concluded that the magnocellular division of the medial genicculate body is a transitional structure between the posterior ventral nucleus and the parvocellular division of the medial geniculate body, and that in addition, it is connected more closely with the auditory than with the somatosensory system. It is suggested that the somatosensory input into the magnocellular division of the medial geniculate body is formed mainly by fibers of the medial lemniscus.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 2, pp. 133–141, March–April, 1978.     

Electrophysiological characteristics of representation of auditory and somatosensory systems in the turtle midbrain

Experiments on waking curarized turtles showed that auditory representation is located in the mediodorsal zone of the tegmentum, in the torus semicircularis, which contains monomodal auditory and bimodal somatoauditory neurons. The somatosensory system is represented more widely and overlaps in the medial zones with the auditory system. Its focus is located in the lateral zones of the dorsal tegmentum (n. intercollicularis), where monomodal somatic neurons were found. Predominance of contralateral somatic projections was discovered. Frequency-threshold curves, obtained by analysis of evoked potentials, were flattened Y-shaped. The range of frequencies received was 40–6000 Hz and the range of optimal frequencies 100–400 Hz. Responses of midbrain auditory neurons could be divided into three principal types: phasic, tonic, and bursting. Neurons with a phasic type of response were characterized by tuning to one optimal frequency, whereas most neurons with responses of tonic type were equally sensitive to two or even three frequencies.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 14, No. 3, pp. 260–269, May–June, 1982.     

Evoked responses of the lateral geniculate body to photic stimulation in intact and visually deprived rabbits

Evoked responses of the lateral geniculate body and visual cortex to stimulation of the contralateral eye with flashes were recorded in rabbits kept in total darkness from birth until the age of 2–3 months. The experiments were carried out under pentobarbital anesthesia and dark adaptation conditions. By contrast with the marked decrease in amplitude of primary visual cortical responses in the visually deprived animals, the amplitudes and temporal parameters of the LGB evoked responses were virtually identical with the corresponding parameters in the control animals. These results suggest that disturbances arising in rabbits as a result of early visual deprivation are localized mainly at the cortical level. The possible mechanisms lying at the basis of the functional disturbances in rabbits reared under conditions of visual deprivation are discussed.Brain Institute, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 10, No. 5, pp. 504–509, September–October, 1978.     

Responses of secondary sensomotor cortical neurons to electrodermal and acoustic stimulation in waking cats

Unit responses in the second somatosensory cortical projection area (SII) to clicks and electric shocks applied to the contralateral limb were investigated in chronic experiments on cats. In response to specific stimulation for the cortical region studied the discharge frequency of 75% of neurons increased, spontaneous activity of 18% was reduced in frequency or the discharges ceased altogether, and 25% of cells did not respond. In response to "nonspecific" stimulation (clicks) 30% of neurons were activated; the discharge of 25% of cells was inhibited and 45% did not respond. The results of investigation of intersensory convergence of stimuli from different sensory systems showed that a high proportion (55%) of SII neurons give bimodal responses. Another 18% of neurons give a specific response to both adequate and inadequate stimulation. It is suggested that the presence of polysensory convergence of SII neurons and of short pathways for the conduction of sensory information, and also the ability of neurons to acquire polysensory properties during stimulus presentation are evidence of the important role of this cortical region in conditioning.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 9, No. 5, pp. 453–459, September–October, 1977.     

Interaction between responses evoked by acoustic and somatosensory stimuli in neurons of the magnocellular part of the medial geniculate body

Interaction between responses to acoustic clicks and to electrodermal stimulation of the contralateral forelimb was investigated in 78 neurons in the magnocellular part of the medial geniculate body of curarized cats. Of this number, 33 neurons responded by discharges both to clicks and to electrodermal stimulation, 25 responded to clicks only, and 20 to electrodermal stimulation only, or to stimulation of the dorsal funiculus of the spinal cord. Conditioning stimulation evoked inhibition of the response to the testing stimulus in 32 of 33 neurons responding by spike discharges to both clicks and electrodermal stimulation. Electrodermal stimulation inhibited responses to clicks in all the neurons tested which responded only to clicks, whereas clicks evoked inhibition of responses to electrodermal stimulation (or to stimulation of the dorsal funiculus) in only four of the 20 neurons which responded to these types of stimulation only. It is suggested that inhibition of excitability arising in neurons of the magnocellular part of the medial geniculate body during interaction between auditory and somatosensory afferent volleys is based on postsynaptic inhibition.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 4, pp. 368–374, July–August, 1980.     

Comparative analysis of the role of the second and first somatosensory cortical areas in somatic responses of the medullary reticular formation in rats

Evoked potentials and unit activity in the medullary reticular formation were investigated in unanesthetized, curarized rats during cold blocking or after extirpation of the cortical representation of one of the stimulated limbs. Local cooling or extirpation in area SII, unlike blocking of area SI, leads to a small (up to 30%) decrease in amplitude and a very small change (up to 10 msec) in the temporal parameters of evoked potentials arising in the reticular formation in response to electrodermal stimulation of the contralateral limb, whose representation in the cortex was blocked. Predominance of corticofugal influences from SI over those from SII was discovered both in experiments with evoked potentials and during analysis of somatic spike responses of reticular formation neurons. Corticofugal control over activity of the medullary reticular formation in rats exerted by neuronal mechanisms of somatosensory areas SII and SI thus differs both qualitatively and quantitatively.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 15, No. 1, pp. 42–49, January–February, 1983.     

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.     

Unit responses of the somatosensory cortex of the waking cat to stimulation of the ventro-posterolateral thalamic nucleus

Unit responses in the primary somatosensory projection cortex to stimulation of the ventro-posterolateral thalamic nucleus were investigated by extra- and intracellular recording in chronic experiments on cats. Short-latency spike responses of 71.3% of recorded neurons appeared after not more than 4 msec. It is concluded that activation of neurons in this area of the cortex is chiefly monosynaptic and disynaptic. Besides participating in the initial response to the stimulus, one-quarter of the neurons generated after-discharges 120–314 msec later. These after-discharges are based on rebound after IPSPs and additional synaptic activation. Initial inhibition may appear 1.5 msec after stimulation of the ventro-posterolateral nucleus, evidence against the participation of recurrent collaterals in the formation of these IPSPs.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 4, pp. 348–354, July–August, 1973.     

Responses of parietal associative neurons to stimulation of primary sensory areas

Responses of 251 neurons in the anterior part of the middle suprasylvian gyrus to stimulation of primary sensory (auditory, visual, somatosensory) areas and also to acoustic, visual, and somatosensory stimuli were studied in acute experiments on cats anesthetized with chloralose (40 mg/kg) and pentobarbital (20 mg/kg). Three groups of neurons were distinguished by their responses to stimulation of the primary sensory areas: those responding by an increased firing rate (117) or by inhibition (35) and those not responding (99). Responses of 193 neurons to stimulation of the peripheral afferent systems were analyzed. Neurons of the parietal associative cortex responded more frequently to cortical stimulation than to peripheral. By the duration of the latent period of their response to cortical stimulation the neurons were divided into three groups: those with short (less than 20 msec), medium (20–30 msec), and long latent periods (over 30 msec). The first group was the largest.Kemerovo State Medical Institute. Translated from Neirofiziologiya, Vol. 4, No. 5, pp. 524–530, September–October, 1972.     

Responses of upper cervical spinal neurons in cats to stimulation of the brain-stem locomotor region at different frequencies

Synaptic responses of neurons in segments C₂ and C₃ to stimulation of locomotor points in the medulla or midbrain were recorded extracellularly in mesencephalic cats. Neurons generating responses with an index of 0.4–0.6 to stimulation with a frequency of 2 Hz maintained this same index at frequencies of 20–60 Hz. The discharge index of many neurons during stimulation at 2 Hz was low, and it increased to 0.4–0.6 when high-frequency stimulation was used. More than half of the cells were excited by stimulation of both ipsilateral and contralateral locomotor points; one-quarter of the neurons responded to stimulation of locomotor points in both medulla and midbrain. The cells studied were located 1.8–4.2 mm from the dorsal surface of the spinal cord. The mean latencies of responses with an index of not less than 0.5 lay within the range 2–30 msec, with a mode of 2–8 msec. Considerable fluctuations of latent period were observed for long-latency responses. The possibility that the neurons studied may participate in the transmission of activity from the locomotor region of the brain stem to stepping generators in the spinal cord is discussed.Institute for Problems of Information Transmission, Academy of Sciences of the USSR, Moscow. M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 15, No. 4, pp. 355–361, July–August, 1983.     

Psychophysiological Features of Somatosensory Disorders in Victims of the Chernobyl Accident

Participants of the Chernobyl clean-up (n = 145) teams exposed to radiation doses from 0.05 to 3.5 Gy who had for the first time complained of pathologic somatosensory sensations (ostealgic syndrome), 20 healthy subjects, and 50 veterans of the war in Afghanistan with posttraumatic stress disorder (PTSD) were examined by a neuropsychiatrist and presented with the MMPI test. Somatosensory evoked potentials (SSEPs) were recorded. Paresthesia and cenesthopathy were characteristic of the participants of the Chernobyl clean-up. Sensation disorders of the cerebral type, kinesthetic illusions, cenesthopathic hypochondriac disorders, and paroxysmal psychosensory states predominated in this group of subjects. They differed significantly from the veterans with PTSD in markedly increased scores on MMPI scales (hypochondriasis, schizophrenia, pure hypochondriasis, pure schizophrenia, emotional exclusion, and perception oddity), which closely correlated with clinical somatosensory symptoms. In clean-up workers, somatosensory disorders were significantly associated with hypochondriac and schizophrenic-like symptoms. The latencies (LPs) of main SSEP components—N ₂₀, P ₂₅, N ₁₄₀, P ₃₀₀, and N ₄₀₀—were increased and their amplitudes decreased in subjects exposed to radiation. Their SSEPs had significant topographical deviations in the left temporoparietal area: the contralateral LPs were increased, whereas the contralateral amplitudes of the thalamocortical N ₂₀ component and the cortical P ₂₅ component were decreased as compared to normal values. Somatosensory disorders and hypochondriac and schizophrenic symptoms were significantly correlated with changes in the SSEPs. The decrease in the N ₂₀ amplitude and increase in the P ₂₅ latency in the left temporoparietal area were dose-dependent. The results suggest cerebral rather than peripheral origin of ostealgic syndrome and other somatosensory disorders in the participants of the Chernobyl clean-up. These disorders are associated with radiation-induced dysfunction of the corticolimbic structures of the left—dominant—hemisphere. It is suggested that somatosensory disorders in patients exposed to low doses of radiation can be considered as manifestations of chronic fatigue syndrome /fibromyalgia, whereas schizoform organic brain lesions manifest themselves after exposure to a radiation dose of 0.3–0.5 Gy.     

Unit responses of the cat somatosensory cortex to stimulation of the reticular and anteroventral thalamic nuclei

Responses of 189 neurons of the somatosensory cortex to stimulation of the nonspecific reticular (R) and anteroventral (AV) nuclei of the thalamus were studied in cats anesthetized with thiopental and immobilized with tubocurarine. In the series of experiments with stimulation of R and, for comparison, of the specific ventral posterolateral nucleus (VPL), 132 neurons were recorded, of which 22 (16.7%) did not respond to stimulation of these nuclei, 77 (58.3%) responded only to stimulation of VPL, and 33 (25%) responded to stimulation of both VPL and R. In the series of experiments in which AV was stimulated, 57 neurons were recorded. Eight (14.8%) responded to neither stimulus and 25 (43.1%) responded only to stimulation of VPL; 24 responded to stimulation of AV (42.1%), and of these, 10 also responded to stimulation of VPL. A characteristic feature of unit responses in the somatosensory cortex to stimulation of the nonspecific nuclei was the irregularity of the responses and their longer latent period. Only five cells responded sooner to stimulation of the nonspecific nuclei than to stimulation of VPL. Responses of the nonspecific nuclei to stimulation appeared clearly only if the stimulation was repetitive. Preliminary stimulation of R blocks the response to stimulation of VPL during the subsequent 40–60 msec.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol.4, No.4, pp. 384–390, July–August, 1972.     

Unit responses of the secondary somatosensory cortex during defensive conditioning in cats

Unit responses in the secondary somatosensory cortex during the formation and extinction of a defensive conditioned reflex to acoustic stimulation were investigated in chronic experiments on cats. In 21 of 28 neurons tested during defensive conditioning the firing pattern changed in accordance with the character of responses to electric shock reinforcement. Two types of conditioned-reflex unit responses were distinguished: excitatory and inhibitory. Most neurons responding to the conditioned stimulus by activation did so during the first 50 msec, which was 80–100 msec before the conditioned motor response. Considerable variability of the unit responses was observed during conditioning. By the time of stabilization of the conditioned-reflex connections the unit response to the conditioned stimulus was stable in form. The pattern of extinction of the conditioned unit activity was expressed as a decrease in the discharge frequency in responses of excitatory type and disinhibition of activity in the case of inhibitory responses.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev, Translated from Neirofiziologiya, Vol. 9, No. 3, pp. 232–238, May–June, 1977.     

Comparison of spontaneous cortical unit activity in several brain areas of unanesthetized cats

Spontaneous unit activity in association area 5 and some projection areas of the cortex (first somatosensory, first and second auditory areas) were studied in cats immobilized with D-tubocurarine in which the index of specific spontaneous activity, the mean frequency, types of spontaneous activity, and statistical parameters — distribution of interspike intervals and autocorrelation function — were determined. The results showed that spontaneous unit activity in the association area differs from that in the projection areas in both intensity and character. A special feature of the spontaneous activity of the auditory areas was a well-marked volley distribution of activity. In the somatosensory area the level of spontaneous activity as reflected in all indices was the lowest. In the association cortex the largest number of neurons with spontaneous activity lay at a depth of 500–1000 µ corresponding to cortical layers III–IV. In the first auditory area neurons with spontaneous activity were concentrated at a depth of 1400 µ (layer V) and in the somatosensory area at a depth of 1000–1400 µ (alyers IV–V). The possible functional significance of these differences is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 1, pp. 13–21, January–February, 1976.     

Comparative effect of thiopental sodium on cortical evoked potentials of different origin

The minimal doses of thiopental sodium (TS) to suppress each of seven varieties of cortical evoked potentials tested were determined in experiments on cats. The preparation had the strongest effect on the following responses: interzonal, arising in the motor cortex to stimulation of somatosensory area I; the reflex pyramidal and late components of the direct pyramidal response. The oligosynaptic transcallosal potential is less sensitive than the above responses but more so than the callosal response. The action of TS on the response depends on the complexity of its organization. The level at which the given cortical response takes place must also be considered. Comparison of the action of TS on the I₁ wave of the pyramidal response and on the dendritic component of the direct cortical response showed that, given equal complexity of its organization, the response arising in deep layers of the cortex is depressed more by TS than the response arising wholly at the level of surface layer I.Institute of Pharmacology, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 4, No. 3, pp. 264–271, May–June, 1972.     

Tegmento-thalamic relations in turtles

A focus of evoked potentials and the largest number of spike responses of single units to tegmental stimulation were found in immobilized turtles (Emys orbicularis), lightly anesthetized with chloralose, in the ventral region of the thalamus, which is a heterosensory zone of predominantly somatosensory modality. Incomplete separation of lateral- and medial-tegmental projections between the dorsal and ventral portions of the ventral thalamic region respectively was discovered. Tegmental stimulation, coinciding with or preceding electrodermal or photic stimulation, causes predominant depression of somatic single unit responses, but has less effect on visual responses. The possibility of comparison of tegmento-thalamic systems in reptiles and mammals is discussed.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 12, No. 2, pp. 165–174, March–April, 1980.     

Responses of somatic cortical neurons during the conditioned placing reflex in cats

Unit activity was studied in areas 3 and 4 during the conditioned placing reflex in cats. Responses of somatic cortical neurons in this case were shown to develop comparatively late — 80–100 or, more often, 200–450 msec after the conditioned stimulus. In the motor cortex responses preceded movement by 50–550 msec, whereas in the somatosensory cortex they usually began simultaneously with or after the beginning of the movement. Judging from responses of somatic cortical neurons, the placing reflex is realized by the same neuronal mechanism as the corresponding voluntary movement. The differential stimulus and positive conditioned stimulus, after extinction of the conditioned placing reflex, evoked short-latency spike responses lasting 250–350 msec in the same neurons as took part in the reflex itself. In these types of internal inhibition, responses of the neurons were thus initially excitatory in character. Participation of the neurons in the conditioned placing reflex and its extinction, disinhibition, and differentiation, is the result of a change in the time course of excitatory processes and is evidently connected with differential changes in the efficiency of the various synaptic inputs of the neuron.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 14, No. 4, pp. 392–401, July–August, 1982.