Responses of medullary neurons to microstimulation of the "locomotor strip" in cats

Synaptic responses of medullary neurons to stimulation of the bulbar locomotor strip with a current of about 20 µA were studied by an extracellular recording method in mesencephalic cerebellectomized cats. The mean latent period of response of 177 neurons was 3.2 msec. Neurons in which synaptic responses appeared were located in both the lateral and the medial parts of the reticular formation, but short-latency responses were recorded predominantly in the lateral part. In response to a single stimulus 32% of neurons generated a discharge of 2–4 spikes. "Respiratory" neurons were not excited by stimulation of the locomotor point. The results indicate that neurons of the locomotor strip may have an excitatory action not only on each other, but also on neurons located medially. The possible mechanisms of the spread of activity to the superior cervical segments of the spinal cord are discussed.Institute for Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 13, No. 3, pp. 275–282, May–June, 1981.     

Unit responses in the "locomotor strip" of the cat hindbrain to microstimulation

Synaptic responses of single units in the "locomotor strip" of the hindbrain were recorded extracellularly. Short-latency responses appeared in neurons of the rostral part of the strip to stimulation of the "locomotor region" of the mesencephalon. Neurons of the caudal part of the strip responded to microstimulation of its other regions, including rostral. If the distance between the neuron and point of stimulation was under 2–3 mm, short-latency (1.2–1.6 msec) responses could be observed. The thresholds and latent periods of the responses increased when the distance apart increased. Polysynaptic responses with a latent period of 3–4 msec could be potentiated by an increase in the frequency of stimulation up to 30–40 Hz. It is suggested that axons of the "locomotor strip" are oriented in the rostrocaudal direction for a distance of 2–3 mm and give off collaterals which run toward neighboring neurons. The strip may be an integrative center, "intercalated" between the rostral portions of the brain stem and spinal cord.Deceased.Institute for Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 10, No. 5, pp. 510–518, September–October, 1978.     

Synaptic responses of propriospinal neurons to stimulation of the stepping strip of the dorsolateral funiculus in cats

Synpatic responses were recorded extracellularly from single neurons at levels T12–T13 in response to microstimulation of the stepping strip of the dorsolateral funiculus in the thoracic and cervical portions of the spinal cord in cats decerebrated at the precollicular level [4]. The latent periods of these responses increased when the distance between the stepping point and recording point exceeded 20 mm, and when two stimuli had to be applied in order to evoke responses. Axons of neurons respondingtrans-synaptically to stimulation of the stepping strip were located on the boundary between the lateral and ventral funiculi close to the gray matter. Antidromic responses of neurons sending their axons in the caudal direction were found on average in 37% of cases, provided that the distance between the recording and stimulation points did not exceed 40 mm. Half of the neurons which generated synaptic responses to stimulation of spinal stepping points could be excited also by a series of three to six stimuli, applied to the mesencephalic locomotor point [15]. It is suggested that the neurons discovered may participate in the spread of activity toward the generator of stepping movements of the hind limb.Institute for Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 17, No. 2, pp. 270–278, March–April, 1985.     

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.     

Activity of propriospinal neurons in segments C3 and C4 during fictitious locomotion in cats

Activity of propriospinal neurons in segments C3 and C4 was recorded in immobilized decerebrate cats, whose spinal cord was divided at the lower thoracic level, during locomotor activity of neuronal mechanisms controlling the forelimbs (fictitious locomotion of the forelimbs). Neurons were identified according to antidromic responses to stimulation of the lateral column of the spinal cord at level C6. Antidromic responses also appeared in 70% of these neurons to stimulation of the medullary lateral reticular nucleus. During fictitious locomotion, i.e., in the absence of afferent signals from the limb receptors, rhythmic modulation of the discharge of most neurons was observed, correlating with activity of motoneurons. If the rostral region of the cervical enlargement of the spinal cord was cooled, causing generation of the locomotor rhythm to cease, rhythmic activity of propriospinal neurons in segments C3 and C4 also ceased. The main source of modulation of activity of propriospinal neurons in segments C3 and C4 is thus the central spinal mechanisms controlling activity of the forelimbs.Institute for Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. M. V. Lomonosov Moscow University. Translated from Neirofiziologiya, Vol. 17, No. 3, pp. 320–326, May–June, 1985.     

The rhombencephalic "locomotor region" in turtles

Electrical stimulation (10–20 µA, 20–30 Hz) of the rhombencephalon in decerebrate turtles can induce cyclic coordinated limb movemnts. The "locomotor region" is a strip, oriented in the rostro-caudal direction, which coincides in its location with the lateral reticular formation. Both in the medial and in the lateral reticular formation extracellular ipsilateral and contralateral synaptic responses of single neurons evoked by stimulation of the "locomotor region," (10–30 µA, 2 Hz), were recorded. Usually these responses had latent periods of between 3 and 12 msec (mode 5–6 msec). Excitation of the "locomotor region" thus leads to extensive spread of activity in the rhombencephalon. The possible mechanisms of this spread are discussed.Institute of Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 12, No. 4, pp. 382–390, July–August, 1980.     

Responses of bulbar neurons to stimulation of locomotor and inhibitory sites in the cat brainstem

Bulbar locomotor and inhibitory sites were located in the pons of mesencephalic decerebellate cats. Rhythmic stimulation of locomotor sites through microelectrodes at the rate of 60 Hz elicited stepping movements in the forelimbs which were halted when the inhibitory sites were rhythmically stimulated. Neuronal response was elicited by single or paired stimulation of locomotor sites at the rate of 1.5 Hz or by applying a series of 2–4 stimuli spaced 2 msec apart to the inhibitory site. Medial neurons generated synaptic responses (postsynaptic potentials or action potentials) to stimulation of the inhibitory site twice as frequently as when the locomotor site was stimulated. Responses in lateral neurons, however, occurred twice as frequently to stimulation of the locomotor site, while IPSP were only observed half as often as EPSP in neurons of both groups. In neurons excited by stimulation of the locomotor site, stimulation of the inhibitory site did not normally produce IPSP. Possible mechanisms underlying the halt of locomotion occurring in response to stimulation of the inhibitory site are discussed.Information Transmission Institute, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 18, No. 4, pp. 525–533, July–August, 1986.     

Propagation of activity along the “stepping strip” of the cat spinal cord

Three points located approximately 8 mm apart were identified in a dorsolateral funiculus of the lower thoracic spinal cord in mesencephalic cats, each producing stepping movements on the ipsilateral hindlimb when stimulated. An area 5–17 mm caudal to the caudal stepping point (SP) was scanned for neurons responding synaptically to stimulating the rostral or caudal SP prior and subsequent to electrolytic coagulation of the medial SP. Relative incidence of neurons excited by stimulating the caudal SP did not change following this type of lesioning, although stimulation of the rostral SP at the rate of 4 Hz induced response 5 times less frequently than before. Even stimulation of the rostral SP at the rate of 40–60 Hz, which had considerably increased firing index prior to coagulation, could only produce excitation in tiny numbers of neurons. This indicates that synaptic excitation of neurons becomes considerably more difficult once the stepping strip between stimulation and recording sites has been damaged.Institute for Research into Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 20, No. 6, pp. 763–769, November–December, 1988.     

Synaptic processes in neurons of the cervical enlargement of the cat spinal cord during stimulation of propriospinal pathways of the dorsolateral tract

Responses of motoneurons and interneurons of the cervical enlargement of the cat spinal cord were studied by a microelectrode technique during selective stimulation of propriospinal fibers of the dorsolateral tract of the lateral white column. The long descending and ascending pathways were blocked by preliminary (10–16 days earlier) hemisection of the spinal cord cranially and caudally to the segments studied. Stimulation of the dorsolateral tract at a distance of 15–25 mm from the site of recording evoked complex postsynaptic potentials consisting of several successive waves in the motoneurons. The character of the PSPs was not clearly linked with the function of the motoneurons. By their latent periods the components of the PSPs could be placed in three groups. The "primary" components were reproduced in response to stimulation at 50–100/sec whereas the "secondary" and "tertiary" components were weakened or blocked. It is postulated that the "primary" components are evoked through monosynaptic connections between propriospinal fibers of the dorsolateral tract and motoneurons of the forelimb muscles, while the late components are evoked through polysynaptic pathways, including segmental interneurons. Many of these interneurons, located in the ventral horn and intermediate zone, were strongly excited during stimulation of the dorsolateral tract.A. A. Bogomolets' Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 1, pp. 61–69, January–February, 1973.     

Unit responses in the auditory cortex to stimulation of geniculocortical fibers

Extracellular and intracellular single unit responses of neurons of the auditory cortex to electrical stimulation of geniculocortical fibers (GCF) were recorded in experiments on cats immobilized with tubocurarine. The latent period of responses of 15% of neurons to GCF stimulation was 0.3–1.5 msec. It is postulated that they were excited anti-dromically. The latent period of spikes generated by neurons responding to GCF stimulation orthodromically varied from 1.6 to 12 msec. In 28.6% of neurons the latent period was 1.6–2.5 msec. It is postulated that these neurons were excited monosynaptically. Intracellular recording revealed primary IPSPs in response to GCF stimulation in 63.3% of neurons, a brief EPSP followed by a prolonged IPSP in 17.7%, an EPSP-spike-IPSP complex in 12.3%, and subthreshold EPSPs in 7% of neurons. The latent period of the primary IPSPs varied from 1.8 to 11 msec, being 1.8–3.7 in 72%, 3.8–5.7 in 20.0%, and 5.8–11 msec in 8.0% of neurons. The latent period of responses beginning with an EPSP was 1–4 msec (mean 1.8 msec). Orthodromic responses arising 3–10 msec after the antidromic response, and consisting of 3–5 spikes, were recorded in some antidromically excited neurons. Hypotheses regarding the functional organization of the auditory cortex and mechanisms of inhibition in its neurons are put forward on the basis of the results obtained.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 3, pp. 227–235, May–June, 1972.     

Responses of rabbit visual cortical neurons to intracortical electrical stimulation

Responses of rabbit visual cortical neurons to single and repetitive intracortical electrical stimulation were investigated. The stimulating electrode was located 0.7–1.2 mm away from the recording electrode. Response thresholds to single stimulation were as a rule 150–180 µA, whereas to series of stimuli they were 30–60 µA. The latent period to the first spike averaged 5–15 msec but the probability of the initial discharge was very low, namely 3–6%. With an increase in current intensity the duration of the initial inhibitory pause was increased in half of the neurons responding to it, whereas in the rest it was unchanged. After presentation of series of stimuli spontaneous activity was enhanced for a short time (4–6 sec). In about half of the cells the same kinds of discharge dynamics were observed in response to repetitive stimulation (frequency 0.25 Hz) as in responses to light, but more neurons with sensitization of discharge and fewer "habituating" neurons took part in responses to electrical stimulation. It is postulated that stimulation of a given point of the visual cortex evokes excitation of a local neuron hypercolumn and inhibition of neighboring cell columns.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 15, No. 4, pp. 412–419, July–August, 1983.     

Connections of the subthalamic and mesencephalic "locomotor regions" in rats

Responses of single neurons to stimulation of the subthalamic "locomotor region" were recorded extracellularly in the tegmentum mesencephali of rats. Latent periods of response of different neurons varied from 1 to 13 msec. The thresholds of the unit responses usually did not exceed the threshold for eliciting locomotion. The results are evidence of direct and oligosynaptic connections between the "locomotor regions" of the subthalamus and mesencephalon.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 9, No. 3, pp. 275–280, May–June, 1977.     

Two patterns of bulbar neuronal response to microstimulation of locomotor and inhibitory brain stem sites

Synaptic responses (postsynaptic potentials and action potentials) were evoked in mesencephalic decerebellated cats by stimulating pontine bulbar locomotor and inhibitory sites (LS and IS, respectively) with a current of not more than 20 µA in "medial" and "lateral" neurons of the medulla. Some neurons even produced a response to presentation of single (actually low — 2–5 Hz — frequency) stimuli. The remaining cells responded to stimulation at a steady rate of 30–60 Hz only. Both groups of medial neurons were more receptive to input from LS. Lateral neurons responding to even single stimuli reacted more commonly to input from LS and those responding to steady stimulation only to input from IS. Many neurons with background activity (whether lateral or medial) produced no stimulus-bound response, but rhythmic stimulation either intensified or inhibited such activity. This response occurs most commonly with LS stimulation. Partial redistribution of target neurons in step with increasing rate of presynaptic input may play a major part in control of motor activity.Institute for Research into Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 22, No. 2, pp. 257–266, March–April, 1990.     

Functional properties distinguishing individual auditory cortical neurons

Responses of 246 auditory cortical neurons to paired and repetitive stimulation of geniculo-cortical fibers were studied in experiments on cats immobilized with tubocurarine. The refractory period (RP) varied from 1 to 200 msec in different neurons. For neurons excited antidromically it varied from 1 to 3 msec. Among neurons excited monosynaptically there were some with a short (1.3–6 msec), medium, (8–16 msec) or long (30–100 msec) refractory period. Most neurons excited polysynaptically had a RP of mean length. RPs 30–200 msec in length were due to inhibition arising in the neuron after conditioning stimulation. In some neurons, after a short (1.5–2.0 msec) initial period of refractoriness there was a temporary (for 2–3 msec) recovery of responsiveness, followed by another period of ineffectiveness of the testing stimulus lasting 30–100 msec. Barbiturates selectively inhibited long-latency unit responses in the auditory cortex and during their action the number of responding neurons with a mean RP decreased sharply. The results demonstrate functional heterogeneity of auditory cortical neurons responding to an incoming volley of afferent impulses.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 3, pp. 236–245, May–June, 1973.     

Synaptic activation of thoracic interneurons by reticulospinal fibers of the lateral funiculus

Synaptic responses of different functional groups of interneurons in segments T10 and T11 to stimulation of the ipsilateral and contralateral medullary reticular formation were investigated in anesthetized cats with only the ipsilateral lateral funiculus remaining intact. Activation of reticulospinal fibers of the lateral funiculus with conduction velocities of 30–100 m/sec was shown to induce short-latency and, in particular, monosynptic EPSPs in all types of cells tested: in interneurons excited by group Ia muscle afferents, in cells activated only by high-threshold cutaneous and muscle afferents (afferents of the flexor reflex), in cells activated mainly by descending systems, and, to a lesser degree, in neurons connected with low-threshold cutaneous afferents. These cell populations are located mainly in the central and lateral parts of Rexed's lamina VII. Most neurons in laminae I–V of the dorsal horn, except six cells located in the superficial layers of the dorsal horn, received no reticulofugal influences. The functional organization of connections of the lateral reticulospinal tract with spinal neurons is discussed and compared with the analogous organization of the medial reticulospinal tract, and also of the "lateral" (cortico- and rubrospinal) descending systems.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 2, pp. 150–161, March–April, 1978.     

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.     

Electrophysiological investigation of pathways in the middle cervical sympathetic ganglion of the turtle

Single unit responses in the middle cervical sympathetic ganglion ofEmys orbicularis to stimulation of other nerves and changes in these responses during the action of sympathetic blocking agents on the ganglion were investigated. The results showed that some fibers of the cervical sympathetic trunk of the turtle are interrupted in this ganglion. Postganglionic fibers pass out of the ganglion and enter the lateral branch and the sympathetic trunk. Other fibers pass through the ganglion without interruption and, together with postganglionic fibers, leave the ganglion in the cervical sympathetic trunk in a cranial direction. The velocity of conduction of excitation along the preganglionic fibers is between 4–3 and 2–1.5 m/sec and along the postganglionic fibers between 4–2.6 and 0.7–0.5 m/sec (fibers of types B₂ and C). Synaptic delay in the fast-conducting fibers averages 6.6 msec. Preganglionic fast-conducting fibers form synaptic contacts on neurons with type B₂ axons, while preganglionic slow-conducting fibers form contacts on neurons with type C axons. Terminals of two preganglionic fibers differing very slightly in their threshold of excitability, and probably constituting the same group, converge on some neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukranian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 1, pp. 83–89, January–February, 1972.     

Functional properties of propriospinal pathways of the dorsolateral tract of the cat spinal cord

The characteristics of conduction of the excitation wave along propriospinal fibers of the dorsolateral tract of the spinal cord were studied in cats anesthetized with pentobarbital. At a preliminary operation, 10–18 days beforehand, lateral hemisection of the spinal cord was performed, cranially in the lumbar division and caudally and cranially in the cervical division to the segments to be studied, leading to degeneration of the long descending and ascending fibers. During stimulation, the dorsolateral tract developed a composite response consisting of a positive-negative wave recorded up to 60–65 mm (4 or 5 segments) from the point of stimulation. The mean conduction velocity of this wave in the lumbar division was 37.9 m/sec compared with 44.5 m/sec in the cervical division. From its properties as a whole this wave can be regarded as the result of excitation of relatively fast-conducting propriospinal fibers of the dorsolateral tract. If the strength of stimulation was increased, late components began to appear in the response. These were evidently connected with excitation of thinner propriospinal fibers and synaptic activation of other other groups of spinal neurons.A. A. Bogomolets' Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 1, pp. 54–60, January–February, 1973.     

Activity of propriospinal neurons during the scratch reflex in cats

Fictitious scratching, i.e., rhythmic activity of hind-limb motoneurons at the characteristic scratching frequency, was evoked by tactile stimulation of the ear in thalamic cats immobilized with flaxedil. Activity of propriospinal neurons in segments C₁, C₂, and T₄–T₇ was recorded extracellularly. The neurons were identified by their antidromic response to stimulation of their axons in segment L₁. Most neurons did not respond to stimulation of the ear. Some neurons, however, were activated during fictitious scratching. Neurons of the cervical segments responded not only to stimulation of the ear, but also to tactile stimulation of the forelimbs and also to passive movements of those limbs. Neurons of the thoracic segments were activated only by stimulation of the ipsilateral ear; these neurons were inhibited by stimulation of the contralateral ear. The role of the propriospinal neurons in the activation of the spinal mechanisms of scratching 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. 9, No. 5, pp. 504–511, September–October, 1977.     

Responses of medial geniculate body neurons to auditory cortical stimulation

Extracellular and intracellular unit responses of thepars principalis of the medial geniculate body to stimulation of the first (AI), second (AII), and third (AIII) auditory cortical areas were studied in cats immobilized with D-tubocurarine. In response to auditory cortical stimulation both antidromic (45–50%) and orthodromic (50–55%) responses occurred in the geniculate neurons. The latent period of the antidromic responses was 0.3–2.5 msec and of the orthodromic 2.0–18.0 msec. Late responses had a latent period of 30–200 msec. Of all neurons responding antidromically to stimulation of AII, 63% responded antidromically to stimulation of AI also, confirming the hypothesis that many of the same neurons of the medial geniculate body have projections into both auditory areas. Orthodromic responses of geniculate neurons consisted either of 1 or 2 spikes or of volleys of 8–12 spikes with a frequency of 300–600/sec. It is suggested that the volleys of spikes were discharges of inhibitory neurons. Intracellular responses were recorded in the form of antidromic spikes, EPSPs, EPSP-spike, EPSP-spike-IPSP, EPSP-IPSP, and primary IPSP. Over 50% of primary IPSP had a latent period of 2.0–4.0 msec. It is suggested that they arose through the participation of inhibitory interneurons located in the medial geniculate body.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 1, pp. 5–12, January–February, 1976.