Analysis of evoked discharges in hypothalamic neurons to somatosensory,acoustic, and photic stimulation

Single unit activity in the supramammillary, mammillary, and anterior hypothalamic areas in response to acoustic, photic, and sciatic nerve stimulation was recorded in cats anesthetized with chloralose and immobilized with succinylcholine. In response to sensory stimulation the spontaneous firing rate was increased or decreased, and silent neurons were activated. Evoked potentials of the silent neurons had the shortest latent period to acoustic and somatosensory stimulation (15 msec), and rather longer to photic stimulation (30 msec); in some cases their latent period was 200 msec. Histograms of interspike interval distribution showed a maximum for intervals of up to 50 msec. Histograms of spike distribution relative to the beginning of stimulation showed maximal density between 100 and 200 msec. A high degree of convergence of excitation was found on units of the anterior as well as the posterior hypothalamus. Unit responses in the hypothalamus to sensory stimuli of all three modalities are regarded as being of secondary, nonspecific type.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 3, No. 6, pp. 592–598, November–December, 1971.     

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.     

Unit activity in the posterior sigmoid and proreal gyri of the cat cortex during bemegride administration

Unit activity was studied in the posterior sigmoid and proreal gyri of the cat cortex in response to photic, acoustic, and somatic stimulation. The animals were first anesthetized with pentobarbital and, after recovery, were given intravenous injections of bemegride (β-ethyl-β-methylglutarimide) and immobilized with diplacin [1, 3-di(β-platyneciniumethoxy)-benzene hydrochloride]. The state of the preparation used was similar to the state of waking animals as regards the characteristics of its unit activity. Unit responses of the proreal gyrus were chiefly modally nonspecific and consisted of brief suppression of spontaneous activity for 100–200 msec. Neurons of the posterior sigmoid gyrus were characterized by much greater modal specificity and variety of types of responses to stimuli of different modalities. Comparison of the characteristics of unit activity in these parts of the neocortex suggests that the frontal cortex in cats is still a highly unspecialized structure.     

Conditions of appearance of off responses of sensomotor cortical neurons to photic stimulation in cats

In acute experiments on cats under chloralose anesthesia (70 mg/kg) unit activity was recorded extra- and intracellularly in the sensomotor cortex (areas 4 and 6) during prolonged (up to 1000 msec) photic stimulation. Responses of on-off type were generated by 100% of neurons tested to photic stimuli whose duration corresponded to the recovery cycle of functional changes after a single flash, determined by the paired stimulation method. Cutaneous stimulation affected the appearance of the photic off response if it led to a spike discharge of the neuron before the off response. It is suggested that IPSPs of cortical neurons largely determine both the duration of the cycle of functional recovery after a single flash and also differences in the pattern of generation of the off response and its interaction with responses to cutaneous stimulation.A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 9, No. 4, pp. 355–360, July–August, 1977.     

Responses of cortical neurons to viscero-somatic stimulation

Responses of cortical neurons in the posterior sigmoid gyrus of cats anesthetized with chloralose to electrical stimulation of somatic and visceral nerves were recorded. Bimodal viscero-somatic neurons are predominant in this part of the cortex and some of them also respond to light. Besides the polysensory modally-specific neurons it was also possible to distinguish a group of modally nonspecific cells (27%), whose responses to different stimuli did not differ statistically from each other. Simultaneous stimulation of visceral and somatic nerves led to facilitation of activity of the long-latency neurons; this was reflected in a decrease of 10 msec in the latent period of the response and an increase in the number of spikes per discharge.A. A. Zhdanov Leningrad University. Tadjik University, Dushanbe. Translated from Neirofiziologiya, Vol. 3, No. 6, pp. 574–581, November–December, 1971.     

Responses of neurons of reticular and ventral anterior nuclei of the cat thalamus to afferent stimuli of different modalities

Responses of 146 spontaneously active neurons of the reticular nucleus (R) and of 98 neurons of the ventral anterior (VA) nucleus of the thalamus to electrical stimulation of the skin of the footpads, to flashes, and to clicks were studied in experiments on cats immobilized with D-tubocurarine or myorelaxin. Stimulation of the contralateral forelimb was the most effective: 24.9% of R neurons and 31.3% of VA neurons responded to this stimulation. A response to clicks was observed in only 4.4% of R neurons and 2.4% of VA neurons. Nearly all responding neurons did so by phasic (one spike or a group of spikes) or tonic excitation. Depression of spontaneous activity was observed only in response to electrical stimulation of the skin. Depending on the site of stimulation, it was observed in 2.6–4.3% of R neurons and 1.7–2.1% of VA neurons tested. The latent period of the phasic responses of most neurons was 6–64 msec to electrical stimulation of the contralateral forelimb, 11–43 msec in response to stimulation of the hindlimb on the same side, 10–60 msec to photic and 8–60 msec to acoustic stimulation. Depending on the character of stimulation, 75.1–95.6% of R neurons and 68.7–97.6% of VA cells did not respond at all to the stimuli used. Of the total number of cells tested against the whole range of stimuli, 25% of R neurons and 47% of VA neurons responded to stimulation of different limbs, whereas 16% of R neurons and 22% of VA cells responded to stimuli of different sensory modalities. The functional role of the convergence revealed in these experiments is to inhibit (or, less frequently, to facilitate) the response of a neuron to a testing stimulus during the 40–70 msec after conditioning stimulation.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 7, No. 6, pp. 563–571, November–December, 1975.     

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.     

Microelectrode analysis of neuronal organization of the parietal association cortex

Responses of 304 neurons in the anterior part of the middle suprasylvian gyrus in cats to acoustic, photic, and somatosensory stimulation and also to simultaneous presentation of 2 or 3 stimuli of these modalities. Three groups of neurons were distinguished: those responding by an increased firing rate (187) or by inhibition (22) and those not responding (95). The first group comprised mono- (64) and polysensory (105) and well as neurons responding only to a combination (18). On the basis of the convergent properties of the polysensory neurons in this region it is postulated that the parietal cortex performs principally integrative processes based on iteraction of visual-cutaneous and cutaneo-auditory afferent information.State Medical Institute, Kemerovo. Translated from Neirofiziologiya, Vol. 4 No. 1, pp. 54–60, January–February, 1972.     

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.     

Unit responses in the posterior suprasylvian gyrus of cats to different stimuli

Extracellular responses of 151 spontaneously active neurons in a small area of the cortex of the posterior suprasylvian gyrus to flashes, clicks, and electrodermal stimulation were studied in unanesthetized cats immobilized with D-tubocurarine. Altogether 63% of neurons responded to the stimuli, of which flashes were the most effective. The proportions of polybi-, and monosensory responding neurons were 60, 18, and 22% respectively. Responding neurons were found throughout the thickness of the cortex, but most frequently at depths of 1000–2000 µ from the brain surface. The latent periods varied not only for different cells (from 20 to 90 msec to all stimuli), but also for the same cell. Responses were unstable, prolonged (over 1 sec) and complex in their dynamic pattern (several phases of increase and decrease in frequency of spontaneous discharges or merely a prolonged increase or decrease in its frequency). In the character of their responses the neurons were divided into 4 groups: 1) poly- and bisensory with equivalent responses to all stimuli; 2) poly- and bi-sensory with nonequivalent responses; 3) monosensory, and 4) nonresponding. The results show that this area of the posterior suprasylvian gyrus is part of the associative cortex with projection predominantly of the visual receptor.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 4, pp. 375–383, July–August, 1972.     

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

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

Neuronal response in nucleus reticularis thalami and neighboring thalamic structures to natural stimulation during chronic experiments on cats

Spike response was investigated in 104 neurons of the nucleus reticularis thalami (R) and adjoining thalamic nuclei to acoustic, tactile, and visual stimuli during chronic experiments on cats. Of the test neurons, 29% responded to acoustic stimulation and 11% showed no preference in relation to different acoustic stimuli. Minimum latencies of response to sounds measured 12–37 msec in excitatory and 18–27 msec in inhibitory cells. Duration of excitation produced by acoustic stimuli reached 50–250 msec; inhibition lasted 27–190 msec. Most cells belonging to this nucleus were excited by different stimuli; the proportion of inhibitory neurons did not exceed 4–10%.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 4, pp. 451–461, July–August, 1989.     

Characteristics of unit responses of the cat cerebellar cortex to acoustic stimuli

Investigation of unit responses of the cerebellar cortex (lobules VI–VII of the vermis) to acoustic stimulation showed that the great majority of neurons responded by a discharge of one spike or a group of spikes with a latent period of 10–40 msec and with a low fluctuation value. Neurons identified as Purkinje cells responded to sound either by inhibition of spontaneous activity or by a "climbing fiber response" with a latent period of 40–60 msec and with a high fluctuation value. In 4 of 80 neurons a prolonged (lasting about 1 sec or more), variable response with a latent period of 225–580 msec was observed. The minimal thresholds of unit responses to acoustic stimuli were distributed within the range from –7 to 77 dB, with a mode from 20 to 50 dB. All the characteristics of the cerebellar unit responses studied were independent of the intensity, duration, and frequency of the sound, like neurons of short-latency type in the inferior colliculi. In certain properties — firing pattern, latent period, and threshold of response — the cerebellar neurons resemble neurons of higher levels of the auditory system: the medial geniculate body and auditory cortex.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 5, No. 1, pp. 3–12, January–February, 1973.     

Postsynaptic potentials of motoneurons evoked by rubrofugal impulsation in the hypoglossal nucleus of the cat

The effect of stimulation of the ipsilateral and contralateral red nuclei on motoneurons of the hypoglossal nucleus was studied in cats anesthetized with chloralose and pentobarbital. In 35 (69%) of the 51 motoneurons tested, PSPs were generated in response to stimulation of the red nuclei by series of 3 to 5 stimuli of threshold strength and with a frequency of 500–600/sec. Of this number, 33 motoneurons responded to stimulation by EPSPs, whose latent periods varied from 3.5 to 14.0 msec (mean value for the ipsilateral red nucleus 5.7±0.75, for the contralateral nucleus 6.8±0.8 msec), whereas two motoneurons responded (after 6.2 msec) by IPSPs. Of the 35 motoneurons responding to stimulation of the red nuclei, stimulation of the lingual nerve evoked EPSPs in 31 and IPSPs in 4 (two of them were inhibited by rubrofugal impulses). IPSPs were generated as a result of stimulation of the lingual nerve in 16 motoneurons which did not respond to rubrofugal impulses.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 1, pp. 62–66, January–February, 1978.     

Vestibular responses of neurons in the anterior suprasylvian gyrus in cats and their interaction with responses to acoustic and facial stimulation

Unit responses in the anterior sigmoid gyrus of cats anesthetized with chloralose (70 mg/kg) to vestibular nerve stimulation and their interaction with responses to acoustic and facial nerve stimulation were investigated. The focus of maximal activity of the vestibular projection was shown to lie a little rostrally to the anterior suprasylvian sulcus. The modality specificity of this part of the cortex to vestibular impulses is reflected in the shortest values of latent periods and the distinct phasic character of the responses, and also in the numerical preponderance of neurons with short-latency responses. Although considerable topographic overlapping of the vestibular projection by acoustic and somatosensory (facial zone) projections is found, the vestibular afferent input predominates over the other afferent inputs.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 13, No. 4, pp. 353–358, July–August, 1981.     

Habituation of postsynaptic unit responses of the cat motor cortex to stimuli of different modalities

Habituation (extinction) of postsynaptic unit responses of the cat motor cortex to repetitive electrodermal, photic, acoustic, and combined bimodal stimulation was investigated by intracellular recording. Habituation was shown by a decrease in the number of spikes per grouped discharge and a decrease in the amplitude and duration of the EPSPs, and sometimes IPSPs, on repetition of the stimulus. The way in which the course of habituation depends on the modality and duration of stimulation (at a constant frequency of 1/sec) is examined. Habituation of postsynaptic responses to sensory stimuli is observed with neurons of different functional groups, namely identified neurons of pyramidal tract and unidentified neurons, some of which were evidently pyramidal neurons and interneurons. The hypothesis is put forward that the habituation of PSPs of the cortical neurons is based on processes taking place mainly at the subcortical level.A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 4, No. 5, pp. 545–553, September–October, 1972.     

Unit responses in the anterior zone of the suprasylvian gyrus to peripheral stimuli of different modalities

Unit responses in the anterior zone of the suprasylvian gyrus to visual, electrodermal, and acoustic stimulation were investigated in experiments on unanesthetized cats immobilized with tubocurarine. Electrical activity was recorded from 131 units, 121 of which were spontaneously active. In 65.5% of cells responses consisted of a short or long increase or a decrease in intensity of spike activity. Most cells (58.2%) were monosensory. Responses to visual stimulation were given by 72% of neurons, to electrodermal by 61.6%, and to acoustic by 9.3%. The corresponding latent periods were 20–40, 20–30, and 15–20 msec. Responses of the same neurons to different peripheral stimuli were uniform or they differed in their dynamics. Intracellular recording gave responses in the form of EPSPs (amplitude 4–5 mV, duration 60–80 msec) or, rarely, IPSPs (amplitude 2–3 mV, duration 160–200 msec). The functional organization of the associative cortex and mechanisms of analysis of incoming afferent information are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 4, pp. 368–374, July–August, 1972.     

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.     

Response of neostriatal neurons to direct electrical stimulation of the optic tract and photic stimulation in awake cats

During acute experiments on awake cats the response of 98 neurons belonging to the head and tail of the caudate nucleus to direct electrical stimulation of the optic tract and presentation of photic stimuli was investigated using extracellular recording techniques. Of the test neurons 34.6% responded to stimulation of the optic tract and 36.2% to optic stimulation. Long latency (over 40 msec for the optic tract and over 80 msec for visual stimulation) excitatory responses prevailed in both cases. A small number of cells responded to optic tract stimulation with short latencies of 5–14 msec. Both types of stimulation were presented during investigations of 58 units of which eight were found to respond to both stimuli. The latter varied in their reaction to different stimuli and their response pattern. Findings are discussed in relation to the possible pathways by which visual information reaches the cortical structure under study.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 18, No. 4, pp. 476–485, July–August, 1986.     

Unit responses in the anterior and posterior hypothalamus to stimulation of the splanchnic and sciatic nerves and to photic stimulation

By extracellular recording of spike discharges the sensory properties of neurons of the anterior and posterior regions of the cat hypothalamus were studied during stimulation of the splanchnic and sciatic nerves and during photic stimulation. Hypothalamic neurons were shown to be characterized by wide convergence of heterosensory excitation: 68% of spontaneously active hypothalamic neurons responded to somatovisceral and photic stimulation. Some posterior hypothalamic neurons responded to somatovisceral stimulation but not to photic stimulation. Neurons responding only to photic stimulation were found in the anterior hypothalamus; no neurons responding only to visceral stimulation were found in the hypothalamus. Total convergence of somatic and visceral afferentation of neurons of the posterior and anterior hypothalamus was observed. Mostly responses of phasic type were obtained to stimulation of all modalities. The study of the quantitative ratio between responses of excitatory and inhibitory types showed that the former predominate. The principles governing the functional organization of hypothalamic afferent systems are discussed.Academician L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 8, No. 3, pp. 276–282, May–June, 1976.