Somatosensory cortical unit responses in the waking cat to afferent stimuli

Extracellular and intracellular responses of 183 neurons in the primary projection area of the somatosensory cortex to electrical and tactile stimulation of the skin on the contralateral fore limb and to stimulation of the ventro-posterolateral thalamic nucleus of the ipsilateral hemisphere were studied in chronic experiments on cats. Spike responses to afferent stimuli are subdivided into three types: initial with a latent period of under 60 msec; initial followed by late responses with a latent period of over 60 msec; late with a latent period of over 60 msec. In addition another group of neurons responding to peripheral stimuli in the interval between the initial and the late response was identified. In nearly all cases the initial responses to peripheral stimulation had the form of a series of spikes, unlike responses to thalamic stimulation. It is concluded from the durations of the latent periods of these responses that about 70% of neurons in the primary projection area are activated mono- and disynaptically in response to peripheral stimulation; consequently, the intracortical spread of excitation in this zone is restricted.     

Neuronal and synaptic mechanisms of cortical inhibition

The latent periods, amplitude, and duration of IPSPs arising in neurons in different parts of the cat cortex in response to afferent stimuli, stimulation of thalamocortical fibers, and intracortical microstimulation are described. The duration of IPSPs evoked in cortical neurons in response to single afferent stimuli varied from 20 to 250 msec (most common frequency 30–60 msec). During intracortical microstimulation of the auditory cortex, IPSPs with a duration of 5–10 msec also appeared. Barbiturates and chloralose increased the duration of the IPSPs to 300–500 msec. The latent period of 73% of IPSPs arising in auditory cortical neurons in response to stimulation of thalamocortical fibers was 1.2 msec longer than the latent period of monosynaptic EPSPs evoked in the same way. It is concluded from these data that inhibition arising in most neurons of cortical projection areas as a result of the arrival of corresponding afferent impulsation is direct afferent inhibition involving the participation of cortical inhibitory interneurons. A mechanism of recurrent inhibition takes part in the development of inhibition in a certain proportion of neurons. IPSPs arise monosynaptically in 2% of cells. A study of responses of cortical neurons to intracortical microstimulation showed that synaptic delay of IPSPs in these cells is 0.3–0.4 msec. The length of axons of inhibitory neurons in layer IV of the auditory cortex reaches 1.5 mm. The velocity of spread of excitation along these axons is 1.6–2.8 msec (mean 2.2 msec).A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 16, No. 3, pp. 394–403, May–June, 1984.     

Vestibular projections in the cat temporal cortex

Boundaries of vestibular projections in the temporal cortex during stimulation of the vestibular nerve were studied in cats anesthetized with pentobarbital and chloralose or chloralose alone. The caudal boundary of the vestibular zone was shown to run along the anterior ectosylvian gyrus. A focus of evoked activity was found in the suprasylvian sulcus or 1–2 mm rostrally to it. All short-latency evoked potentials recorded during vestibular nerve stimulation in the temporal region caudally to the zone mentioned above were connected with the spread of current to auditory structures. To verify the extent of spread of the stimulating current, focal potentials were recorded in the vestibular and superior olivary groups of nuclei. Special experiments were carried out to study the topography of these potentials at the level of bulbar structures during stimulation of vestibular and auditory nerves. According to the results, there is no second vestibular area in the temporal cortex in cats. Vestibular afferentation is projected mainly into the contralateral hemisphere, and the response latency is 5.2±0.7 msec. The ipsilateral evoked potentials had a long latent period (8.4±1.3 msec), and their amplitude depended on the type of anesthesia; it was accordingly postulated that additional synaptic relays exist in this vestibulocortical pathway.     

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.     

Corticofugal influences on the motoneurons of the trigeminal nucleus

We studied the postsynaptic potentials evoked from 76 trigeminal motoneurons by stimulation of the motor (MI) and somatosensory (SI) cortex in the ipsilateral and contralateral hemispheres of the cat. Stimulation of these cortical regions evoked primarily inhibitory postsynaptic potentials (PSP) in the motoneuron of the masseter muscle, but we also observed excitatory PSP and mixed reactions of the EPSP/IPSP type. The average IPSP latent period for the motoneurons of the masseter on stimulation of the ipsilateral cortex was 6.1±0.3 msec, while that on stimulation of the contralateral cortex was 5.2±0.4 msec; the corresponding figures for the EPSP were 7.6±0.5 and 4.5±0.3 msec respectively. Corticofugal impulses evoked only EPSP and action potentials in the motoneurons of the digastric muscle (m. digastricus). The latent period of the EPSP was 7.6 msec when evoked by afferent impulses from the ipsilateral cortex and 5.4 msec when evoked by pulses from the contralateral cortex. The duration of the PSP ranged from 25 to 30 msec. Postsynaptic potentials developed in the motoneurons studied when the cortex was stimulated with a single stimulus. An increase in the number of stimuli in the series led to a rise in the PSP amplitude and a reduction in the latent periods. When the cortex was stimulated with a series of pulses (lasting 1.0 msec), the IPSP were prolonged by appearance of a late slow component. We have hypothesized that activation of the trigeminal motoneurons by corticofugal impulsation is effected through a polysynaptic pathway; each functional group of motoneurons is activated in the same manner by the ipsilateral and contralateral cortex. The excitation of the digastric motoneurons and inhibition of the masseter motoneurons indicates reciprocal cortical control of their activity.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 3, No. 5, pp. 512–519, September–October, 1971.     

Electrophysiological study of connections between the entorhinal cortex and the neocortex

In acute experiments in rabbits immobilized by d-tubocurarine, stimulation of the entorhinal area with rectangular electric impulses led to the appearance of evoked potentials (EP) with a latent period of 6–12 msec in the occipital, temporal, parietal, and cingular areas of the neocortex. The amplitude of the positive response component was 500 µV, and its duration 25–50 msec. The negative component was not always discernible. When rhythmic stimulation was used, these EPs followed stimulation frequencies not exceeding 20 per sec. Stimulation of the medial parts of the entorhinal area with a frequency of one to three per sec was accompanied by recruitment of the EP in the occipital and temporal neocortex areas. Nembutal depressed the amplitude of the neocortex EP appearing in response to stimulation of the entorhinal cortex. With the aid of double stimulation it could be established that, after conditioning stimulation of the entorhinal area, the positive component of the primary response (PR) evoked by stimulation of the contralateral sciatic nerve in the projection zone of the somatosensory cortex is strengthened during the first 50 msec, and subsequently after 80–120 msec. In these cases, the negative component was depressed. These findings are discussed with a view to the influence of limbic structures on the neocortex.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 1, pp. 73–78, January–February, 1970.     

Influence of acute cerebellar lesions on somatosensory evoked potentials (SEPs) in cats.

We studied the effect of acute unilateral cerebellar lesions on the cerebello-thalamo-cortical projection in cats. The lesions were classified into two groups according to their extent. In group I the lesion only covered the cerebellar cortex, while in group II both the cerebellar cortex and deep cerebellar nuclei were removed. Early (short-latency) and late (long-latency) waves, evoked by an electrical stimulation of a forelimb, were collected contralateral to the stimulated leg hemisphere. Pre- and postsurgery recordings from primary and non-primary (motor and parietal) cortices were compared. Cerebellar impairment had a strong influence on discharges of all the considered cortical areas. Early non-primary and primary responses increased in group I and remained unchanged in group II. Late somatosensory evoked potentials components were suppressed in both groups. An inhibitory influence of the cerebellar cortex on the thalamo-cortical projection was confirmed. Changes within the primary sensory cortex may suggest an engagement of that area in the compensation process of cerebellar dysfunction shortly after cerebellar lesion. An alteration in the unaffected hemisphere activation indicate that the spino-cerebellar and cerebello-cortical inputs, responsible for somatosensory evoked potentials generation, are regulated through contralateral and ipsilateral pathways. These pathways are unmasked by cerebellar lesion.     

Functional organization of the kinesthetic projections of the 1st somatosensory region of the cat cortex

Electrophysiological studies were performed on adult cats under ethaminal anesthesia. Kinesthetic potentials were evoked by passive extension of the ulnar joint and recorded in contralateral primary somatosensory cortical area. Natural (nonelectrical) stimulation of peripheral kinesthetic receptors was performed according to the author's original method. The results obtained show significantly shorter latent period of contralateral kinesthetic potentials in comparison with somatosensory potentials in response to electrical stimulation of the skin. These data demonstrate the possibility of super-rapid conduction of modal-specific volleys to the cortical projection centres in the kinesthetic system of cats.     

Somatosensory evoked potentials in healthy people

The results of a study of somatosensory evoked potentials recorded in 37 healthy subjects of both sexes are described. Comparison of the results of tests on three age subgroups showed selectivity in the change in latencies and amplitudes of waves of the somatosensory responses depending on the subject's age. Ipsilateral responses show greater variability but a lower amplitude and frequency of appearance of the individual components than contralateral responses. Their latent periods also were longer than those of the contralateral responses.Institute of Cybernetics, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 5, pp. 447–454, September–October, 1976.     

Distributed current analyses of bi-hemispheric magnetic N1m responses to ipsi/contralateral monaural stimuli from a single subject

Magnetoencephalographic (MEG) responses of both auditory cortices to simple auditory stimuli presented monaurally to either ear were recorded from a single subject. A distributed current model and a current dipole model were used to analyse the responses at the latency of the dominant N1m complex. At the N1m the current density was localised to a single area and was consequently well modelled by a single current dipole close to the peak current density. In the left hemisphere, the contralateral response (as identified by the peak current density) preceded the ipsilateral response by 3 msec. This value was 7 msec for the right hemisphere. Evidence was found in the right hemisphere of a posterior-anterior movement along the sylvian fissure. Also, the left hemisphere N1m sources were all represented more posterior than the right hemisphere N1m sources.     

The effects of neck flexion on cerebral potentials evoked by visual,auditory and somatosensory stimuli and focal brain blood flow in related sensory cortices

Background

A flexed neck posture leads to non-specific activation of the brain. Sensory evoked cerebral potentials and focal brain blood flow have been used to evaluate the activation of the sensory cortex. We investigated the effects of a flexed neck posture on the cerebral potentials evoked by visual, auditory and somatosensory stimuli and focal brain blood flow in the related sensory cortices.

Methods

Twelve healthy young adults received right visual hemi-field, binaural auditory and left median nerve stimuli while sitting with the neck in a resting and flexed (20° flexion) position. Sensory evoked potentials were recorded from the right occipital region, Cz in accordance with the international 10–20 system, and 2 cm posterior from C4, during visual, auditory and somatosensory stimulations. The oxidative-hemoglobin concentration was measured in the respective sensory cortex using near-infrared spectroscopy.

Results

Latencies of the late component of all sensory evoked potentials significantly shortened, and the amplitude of auditory evoked potentials increased when the neck was in a flexed position. Oxidative-hemoglobin concentrations in the left and right visual cortices were higher during visual stimulation in the flexed neck position. The left visual cortex is responsible for receiving the visual information. In addition, oxidative-hemoglobin concentrations in the bilateral auditory cortex during auditory stimulation, and in the right somatosensory cortex during somatosensory stimulation, were higher in the flexed neck position.

Conclusions

Visual, auditory and somatosensory pathways were activated by neck flexion. The sensory cortices were selectively activated, reflecting the modalities in sensory projection to the cerebral cortex and inter-hemispheric connections.     

Postsynaptic potentials of neck muscle motoneurons evoked by horizontal semicircular canal stimulation in cats

Postsynaptic potentials evoked by stimulation of ipsilateral and contralateral horizontal semicircular canals in motoneurons of muscles tilting and turning the head were investigated in acute experiments on cats anesthetized with chloralose and pentobarbital. Stimulation of the ipsilateral canal evoked EPSPs with latent periods varying from 1.8 to 10.0 msec in 25 of these motoneurons and IPSPs with latent periods varying from 1.9 to 3.9 msec in 10 of them. Calculation of the impulse conduction time from the ipsilateral semicircular canal through Deiters' nucleus to the cervical motoneurons indicates that EPSPs with latent periods of under 3.8 msec may be regarded as disynaptic, and those with latent periods of over 3.8 msec as polysynaptic. Stimulation of the contralateral canal evoked EPSPs with latent periods varying from 1.8 to 6.0 msec in 19 motoneurons and IPSPs with latent periods varying from 3.2 to 3.9 msec in two cells. The possible pathways of transmission of these influences and their functional role are discussed.     

Whole-head mapping of middle-latency auditory evoked magnetic fields

We recorded middle-latency auditory evoked magnetic fields from 9 healthy subjects with a 122-channel whole-head SQUID gradiometer. The stimuli were click triplets, 2.5 msec in total duration, delivered alternately to the two ears once every 333 msec. Contralateral clicks elicited P30m responses in 16 and P50m responses in 12 out of 18 hemispheres studied; ipsilateral clicks did so in 7 and 13 hemispheres, respectively. The field patterns were satisfactorily explained by current dipoles in 16 and 4 hemispheres for contra- and ipsilateral P30m, and in 4 and 10 hemispheres for contra- and ipsilateral P50m. The peak latencies of P30m and P50m were not affected by stimulation side. The results show that middle-latency auditory evoked responses receive a strong contribution from auditory cortical structures, and that differences of input latency to cortical auditory areas, evaluated from MLAEF latencies, do not explain the latency differences seen in late auditory evoked fields to contralateral vs. ipsilateral stimulation.     

Neuronal responses in an isolated slab of the cat auditory cortex to intracortical stimulation

Neuronal responses in an isolated slab (area AI) to intracortical pulsed electrical stimulation at the level of layer IV were investigated extracellularly in acute experiments on cats immobilized with D-tubocurarine. Responding neurons were found in all layers of the slab. The character of their distribution by depth in the slab depended on the distance between recording and stimulating electrodes. The latent period of responses of different neurons ranged from 0.8 to 25 msec. With interelectrode distances of 0.5–2 mm most neurons responded mono- and disynaptically. However, responses of many neurons had a latent period of over 4 msec, i.e., they were polysynaptic. This indicates the complex character of interneuronal interactions, even in a limited area of the cortex. After intracortical stimulation no after-discharges with a latent period of over 40 msec could be recorded in the isolated slab of auditory cortex.I. I. Mechnikov Odessa State University. Translated from Neirofiziologiya, Vol. 14, No. 1, pp. 85–93, January–February, 1982.     

Early deflections of cerebral magnetic responses to median nerve stimulation

We have recorded early components of somatosensory evoked magnetic fields with a sensitive 7-channel first-order gradiometer using a wide recording passband (0.05–2000 Hz) and high sampling frequency (8000 Hz). The left median nerve was stimulated at the wrist and responses were recorded over the right hemisphere. The responses typically consisted of a N20m peaking at 18–20 msec, a small P22m peaking at 21–23 msec and a P27m peaking at 29–31 msec. The topography of N20m could be explained by a tangential current dipole in the posterior wall of the central sulcus (probably in area 3b). The equivalent dipoles of P27m were located on average 10 mm antero-medially to the sources of N20m. This suggests that P27m may get a contribution from the anterior wall of the central sulcus. An increase of stimulus repetition rate from 2 to 5 Hz decreased the amplitude of P27m more than that of N20m, which implies that these two deflactions are generated by different neural netwoks.     

Unit responses in the cat auditory cortex to medial geniculate body stimulation

Responses of 98 auditory cortical neurons to electrical stimulation of the medial geniculate body (MGB) were recorded (45 extracellulary, 53 intracellularly) in experiments on cats immobilized with tubocurarine. Responses of the same neurons to clicks were recorded for comparison. Of the total number of neurons, 75 (76%) responded both to MGB stimulation and to clicks, and 23 (24%) to MGB stimulation only. The latent period of extracellularly recorded action potentials of auditory cortical neurons in response to clicks varied from 7 to 28 msec (late responses were disregarded), and that to MGB stimulation varied from 1.5 to 12.5 msec. For EPSPs these values were 8–13 and 1–4 msec respectively. The latent period of IPSPs arising in response to MGB stimulation varied from 2.2 to 6.5 msec; for 34% of neurons it did not exceed 3 msec. The difference between the latent periods of responses to clicks and to MGB stimulation varied for different neurons from 6 to 21 msec. Responses of 11% of neurons to MGB stimulation, recorded intracellularly, consisted of sub-threshold EPSPs, while responses of 23% of neurons began with an EPSP which was either followed by an action potential and subsequent IPSP or was at once cut off by an IPSP; 66% of neurons responded with primary IPSPs. Neurons responding to MGB stimulation by primary IPSPs are distributed irregularly in the depth of the cortex: there are very few in layers III and IV and many more at a depth of 1.6–2 mm. Conversely, excited neurons are predominant in layer III and IV, and they are few in number at a depth of 1.6–2 mm. It is concluded that the afferent volley reaching the auditory cortex induces excitation of some neurons therein and, at the same time, by the principle of reciprocity, induces inhibition of others. This afferent inhibition takes place with the participation of inhibitory interneurons, and in some cells the inhibition is recurrent. The existence of reciprocal relationships between neurons in different layers of the auditory cortex is postulated.A. A. Bogomolets' Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 1, pp. 23–31, January–February, 1972.     

The effect of stimulation of the deep layers of somatosensory zones CI and CII on the activity of motor zone MI of the cat cortex]

The influence of electrical stimulation of deep layers of the somatosensory zones CI and CII on unit responses and intercortical evoked potentials (IEP) in the motor cortical zone MI in projection areas of the anterior contralateral limb was studied in cats anaesthetized with Nembutal and immobilized with diplacine. Latencies of the main IEP components and their different behaviour during repeated stimulation, experimental hypoxia and Nembutal administration suggested the presence of intercortical connections of an oligo- and polysynaptic nature. Only 22% of the MI zone units proved to be responsive to CI and CII stimulation; the latencies of the unit discharges varied from 4.3 to 35 msec. A relatively smaller effectiveness of short-latency inputs from CI and CII to MI was recorded as compared with long latency ones.     

SEP topographies elicited by innocuous and noxious sural nerve stimulation. III. Dipole source localization analysis

The dipole source localization method was used to determine which of the brain areas known to be involved in somatosensation are the best candidate generators of the somatosensory evoked potential evoked by sural nerve stimulation. The ipsilateral central negativity and contralateral frontal positivity which occurred between 58 and 90 msec post stimulus (stable period 1) were best represented by a single source located in the primary somatosensory cortex (SI). The symmetrical central negativity and bilateral frontal positivity which occurred between 92 and 120 msec post stimulus (stable period 2) was best represented by 3 sources. One of these sources was located in SI and the other 2 were located bilaterally in either the frontal operculum or near the second somatosensory cortex (SII). The widespread negativity whose minimum was located in the contralateral fronto-temporal region and which occurred between 135 and 157 msec post stimulus (stable period 3) was also best represented by 3 sources. Two of these sources may be located bilaterally in the hippocampus. We cannot, however, eliminate the possibility that multiple sources in the cortex overlying the hippocampus (e.g., SII and frontal cortex) are responsible for these potentials. At innocuous stimulus levels the third source for stable period 3 was located near the vertex, possibly involving the supplementary motor cortex, whereas at noxious levels this source appears to be located in the cingulate cortex. We were unable to achieve any convincing source localization for the widespread positivity which occurred between 178 and 339 msec post stimulus (stable periods 4–6). Available evidence suggests that more sources were active during this interval than the three we could reliably test under these conditions.     

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 neurons in the vibrissae projection area of the cat somatosensory cortex to afferent activation

Responses of 375 primary somatosensory cortical neurons located in the projection area of the vibrissae to electrical stimulation of the infraorbital nerve and also to adequate stimulation of the vibrissae were investigated in unanesthetized cats immobilized with tubocurarine. Stimulation of the nerve and vibrissae most frequently evoked synaptic responses in the neurons, in the form of a short EPSP followed by an IPSP or, less frequently, as a primary IPSP; during extracellular recordings corresponding changes were observed in spike activity. In response to stimulation of the vibrissae, initial inhibition was found more often than to stimulation of the nerve (in 45 and 16% of neurons respectively). The difference between the minimal values of latent periods of IPSP and EPSP evoked by stimulation of the infraorbital nerve was 0.8 msec in different neurons, and the difference between the mean values 1.4 msec. Directional sensitivity of the cortical neurons was demonstrated (to a change in the direction of deflection of the vibrissae). Neurons located close together could differ in the character of their directional sensitivity during stimulation of the same vibrissae. It is concluded that short-latency inhibition arising in the primary projection area of the cat somatosensory cortex is predominantly afferent and not recurrent. The probable mechanisms of directional sensitivity of the neurons studied are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSSR, Kiev. Translated from Neirofiziologia, Vol. 11, No. 6, pp. 550–559, November, 1979.