Convergence of reticulo-, vestibulo-, and rubrospinal influences on cervical spinal motoneurons in cats

Intracellular recordings were made of synaptic responses of 93 motoneurons in the cervical region of the cat spinal cord to stimulation of the medial longitudinal bundle, the brain-stem reticular formation, the lateral vestibular nucleus of Deiters, and the red nucleus. In response to stimulation of the medial longitudinal bundle and the vestibular nucleus responses in the motoneurons of the distal groups of muscles of the forelimb were predominantly excitatory, whereas in motoneurons of the proximal extensor muscles they were predominantly inhibitory. During stimulation of the red nucleus, excitatory and inhibitory responses were recorded in almost equal numbers of cells regardless of their functional class. Monosynaptic EPSPs appeared in one-fifth of motoneurons in response to stimulation of the medial longitudinal bundle and, in a few cases, to stimulation of the vestibular and red nuclei. Otherwise, during stimulation of these structures polysynaptic responses were recorded in the motoneurons. In 62% of cases postsynaptic potentials arising in response to stimulation of the various suprasegmental structures tested were identical in direction in the same motoneurons. A mutually facilitatory effect was observed during stimulation of different suprasegmental inputs. The results are evidence that interaction between influences of the structures tested takes place largely at the level of spinal interneurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 4, pp. 391–399, July–August, 1978.     

Neurophysiology of the motoneurons of the external oculomotor nucleus in the waking cat

The spontaneous and visually induced activity of abducens motoneurons has been recorded in the alert cat. Motoneurons were identified by their antidromic activation from the ipsilateral abducens nerve. All identified motoneurons appeared related to both the position and velocity of the eye in the horizontal plane, although distributed in a wide range. Neural time constants were also measured, showing a mean value similar to that of the mechanical time constant of the oculomotor plant. According to present results, abducens motoneurons of cats and monkeys are very similar, notwithstanding some differences in their activities during saccadic movements.     

An intracellular study of pudendal afferent inputs onto tail motoneurons in the spinalized cat

Postsynaptic potentials, elicited by stimulation of the sensory pudendal (SPud) and superficial perineal nerves (SPeri) on both sides, were recorded from motoneurons innervating tail muscles in the non-anaesthetized and spinalized cat. The stimulation of SPud and SPeri on both sides predominantly produced excitatory postsynaptic potentials (EPSPs) in all kinds of tail motoneurons (70-95%). The inhibitory postsynaptic potentials (IPSPs) were often observed in motoneurons innervating ventral tail muscles (30-33%). The means of averaged central latencies of EPSPs and IPSPs ranged from 4.3 to 7.3 ms, and from 4.6 to 8.4 ms, respectively. The findings suggests that polysynaptic neuronal pathways from pudendal nerve to tail motoneurons produce tonic activities of all tail muscles to raise the tail in micturation, defecation and sexual movements which are induced by stimulation of pudendal nerves.     

Cortico-pyramidal and cortico-extrapyramidal synaptic influences on lumbar motoneurons in monkeys

Postsynaptic potentials evoked by stimulation of the motor cortex or pyramids before and after acute pyramidotomy were investigated in the lumbar motoneurons of monkeys. In response to activation of fibers of the pyramidal tract monosynaptic EPSPs predominated in motoneurons innervating the distal muscles of the hind limbs. Monosynaptic EPSPs in the motoneurons of the distal muscles had a significantly higher amplitude and could be evoked by weaker stimuli than EPSPs in the motoneurons of the proximal muscles. Cortico-motoneuronal EPSPs in the motoneurons of the distal muscles had a less marked frequency potentiation than EPSPs with monosynaptic segmental delay in the motoneurons of the proximal muscles. Cortico-extrapyramidal synaptic responses appeared in the pyramidotomized monkeys during intensive repetitive stimulation of the motor cortex in motoneurons of both distal and proximal muscles. These effects, transmitted by descending projections of the brain stem, may be responsible for the partial preservation of cortical motor control after pyramidotomy.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 4, No. 6, pp. 587–596, November–December, 1972.     

Measurements of simultaneously recorded spiking activity and local field potentials suggest that spatial selection emerges in the frontal eye field

The frontal eye field (FEF) participates in selecting the location of behaviorally relevant stimuli for guiding attention and eye movements. We simultaneously recorded local field potentials (LFPs) and spiking activity in the FEF of monkeys performing memory-guided saccade and covert visual search tasks. We compared visual latencies and the time course of spatially selective responses in LFPs and spiking activity. Consistent with the view that LFPs represent synaptic input, visual responses appeared first in the LFPs followed by visual responses in the spiking activity. However, spatially selective activity identifying the location of the target in the visual search array appeared in the spikes about 30 ms before it appeared in the LFPs. Because LFPs reflect dendritic input and spikes measure neuronal output in a local brain region, this temporal relationship suggests that spatial selection necessary for attention and eye movements is computed locally in FEF from spatially nonselective inputs.     

Feature-based attention in the frontal eye field and area V4 during visual search

When we search for a target in a crowded visual scene, we often use the distinguishing features of the target, such as color or shape, to guide our attention and eye movements. To investigate the neural mechanisms of feature-based attention, we simultaneously recorded neural responses in the frontal eye field (FEF) and area V4 while monkeys performed a visual search task. The responses of cells in both areas were modulated by feature attention, independent of spatial attention, and the magnitude of response enhancement was inversely correlated with the number of saccades needed to find the target. However, an analysis of the latency of sensory and attentional influences on responses suggested that V4 provides bottom-up sensory information about stimulus features, whereas the FEF provides a top-down attentional bias toward target features that modulates sensory processing in V4 and that could be used to guide the eyes to a searched-for target.     

Interaction between segmental and descending intersegmental reflexes in lumbar motoneurons

The effects of segmental reflexes on descending intersegmental reflexes to stimulation of forelimb afferents were studied in anesthetized cats by recording postsynaptic responses from single motoneurons. Interaction between these influences was found to be reciprocal in character for groups of neurons with primary connections with afferents of the superficial and deep branches of the peroneal nerve and afferents of the nerve to the gastrocnemius muscle. Excitatory postsynaptic responses arising in groups of motoneurons of the peroneal nerve to stimulation of forelimb afferents underwent profound and prolonged inhibition during conditioning stimulation of afferents in the deep and superficial peroneal nerves. Activation of segmental afferents during conditioning stimulation of the gastrocnemius nerve was accompanied by inhibition of excitatory intersegmental responses and deinhibition of inhibitory responses in motoneurons of the gastrocnemius muscle. Segmental inhibition of intersegmental descending impulse activity appeared in the interneuron system of the segmental reflex centers connecting the descending propriospinal tracts with the motoneurons of these centers.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 4, No. 2, pp. 16872-175, March–April, 1972.     

Multiple gate control of the descending statocyst-motor pathway in the crayfish Procambarus clarkii Girard

1. Synaptic responses of uropod motoneurons and interneurons to magnetic field stimulation of the statocyst were studied in a whole animal preparation using intracellular recording and staining techniques to characterize the descending statocyst pathways controlling uropod steering behavior. 2. When the animal was engaged in abdominal postural movement, all uropod motoneurons received sustained excitatory input. Motoneurons which were to be activated during steering behavior showed excitatory responses to the stimulus superimposed on the sustained excitation. In the resting state, they showed weaker responses or no visible responses to the same stimulation. 3. Motoneurons to be suppressed during steering showed inhibitory responses to the stimulus only during abdominal movement. These included both active inhibition as well as disfacilitatory suppression of excitatory input to the motoneurons. 4. Premotor nonspiking interneurons, like motoneurons, showed greater responses to the stimulus during abdominal movement than at rest. Unlike motoneurons, however, they did not always receive sustained input during abdominal movement. 5. Descending axons which responded to statocyst stimulation independent of abdominal movement were found in the 4th and 5th abdominal ganglia. Other axons showed greater responses during abdominal movement than at rest. 6. A number of intersegmental descending interneurons with cell bodies and dendrites in the 4th or 5th ganglion were found to receive excitatory inputs from both the statocyst and the motor system controlling abdominal posture. These responses were found to summate with each other to generate spikes. 7. Statocyst signals are thus transmitted to uropod motoneurons by two types of descending pathways: one whose operation is affected by the abdominal system and the other which operates independently. The former pathway functions by recruiting intersegmental abdominal interneurons and makes stronger connections with motoneurons than the latter.     

Effects of repetitive stimulation of the locus coeruleus on spinal inhibitory responses to suprasegmental stimulation in cats

Effects of repetitive stimulation of the locus coeruleus on spinal responses to activation of cortico-, reticulo-, and vestibulospinal tracts were studied in decerebellate cats anesthetized with chloralose. Descending influences of these structures were assessed from changes in amplitude of extensor and flexor monosynaptic discharges or from the magnitude of postsynaptic potentials recorded from the corresponding motoneurons. Stimulation of the motor cortex or modullary reticular formation as a rule evoked two-component inhibitory responses in extensor motoneurons and excitatory-inhibitory responses in flexor motoneurons. Stimulation of locus coeruleus effectively depressed the amplitude of the late component and, to a lesser degree, that of the early component of inhibition arising after stimulation of the cerebral cortex or reticular formation. During stimulation of the locus coeruleus no marked changes were found in inhibitory responses evoked by vestibulospinal influences in flexor motoneurons, and also in excitatory responses arising after stimulation of the above-mentioned descending pathways in both groups of motoneurons.     

Neuronal activity of the external oculomotor nucleus during saccadic eye movement in the waking cat

The activity of antidromically identified abducens nucleus motoneurons and inter-nuclear neurons has been recorded during saccadic eye movements in the alert cat. The activity of these neurons has been demonstrated to be the sum of a velocity component proportional to eye velocity plus a position component proportional to instantaneous eye position during the movement. Results are discussed in relation to proposed models about the generation of saccadic eye movements.     

Rubrofugal activation of motoneurons of the cat accessory nucleus

Postsynaptic potentials evoked in accessory nerve motoneurons by stimulation of the ipsilateral and contralateral red nuclei were investigated in acute experiments on cats anesthetized with chloralose and pentobarbital. Polysynaptic EPSPs with latent periods of 5.2 to 16 (mean 9.1 ± 0.7) msec and from 5.5 to 18 (mean 10.3 ± 0.9) msec, respectively, appeared in motoneurons of the accessory nerve in response to stimulation of the contralateral and ipsilateral red nuclei. A minimum of two or three stimuli was necessary to produce EPSPs in these motoneurons. In response to single stimulation of the contralateral and ipsilateral red nuclei EPSPs appeared in four motoneurons of the trapezius muscle with latent periods of 2.5 to 5.0 and 3.0 to 5.2 msec, respectively. An increase in the number of stimuli led to action potential generation by motoneurons. The functional role of such activation is discussed.A. A. Bogomolets Institue of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 14, No. 5, pp. 532–536, September–October, 1982.     

Amygdaloid-induced jaw opening and facilitation or inhibition of the trigeminal motoneurons in the rat

1. Repetitive stimulation of the rat'se central amygdaloid (CAm) nucleus induced rhythmic masticatory jaw movements or continuous jaw opening. Both types of jaw movements were accompanied by coincidental activities of the mylohyoid (Myl) nerve. 2. The effects of CAm stimulation were examined on activities of bilateral Myl and masseteric (Mass) nerves or their motoneurons (Myl-Dig and Mass, respectively). 3. CAm stimulation induced contralaterally dominant facilitation of the Myl nerve activity as well as Myl-Dig motoneurons. These facilitatory effects were caused by EPSPs seen in Myl-Dig motoneurons. 4. One third of the Mass motoneurons were inhibited or hyperpolarized by contralateral CAm stimulation, while a few were facilitated and the majority unaffected.     

Role of different types of fibers of the infraorbital nerve in synaptic activation of masseter and digastric motoneurons

Postsynaptic potentials of motoneurons of the masseter and digastric muscles evoked by stimulation of the infraorbital nerve with a strength of between 1 and 10 thresholds were investigated in cats anesthetized with a mixture of chloralose and pentobarbital. Depending on their ability to be activated by low-threshold afferents of this nerve, motoneurons of the masseter were divided into two groups. Stimuli with a strength of 1.2–2.5 times above threshold for the most excitable fibers of the infraorbital nerve evoked short-latency EPSPs in the motoneurons of the first group; a further increase in stimulus strength (3–9 thresholds) led to the appearance of IPSPs with latent periods of 2.8–3.5 msec. Motoneurons of the second group responded to stimulation of the infraorbital nerve with a strength of 3–9 thresholds by IPSPs whose latent periods varied from 6 to 8 msec. Stimuli below 3 thresholds in strength evoked no responses in these motoneurons. Stimulation of the infraorbital nerve with pulses of between 1 and 2 thresholds in strength evoked EPSPs in digastric motoneurons, but an increase in the strength of stimulation led to action potential generation. The presence of many excitatory and inhibitory inputs formed by afferent fibers of different types evidently provides a basis for functional diversity of jaw-opening and jaw-closing reflexes.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 6, pp. 596–603, November–December, 1980.     

Afferent signals from pigeon extraocular muscles modify the vestibular responses of units in the abducens nucleus.

Although the extraocular muscles contain stretch receptors it is generally believed that their afferents exert no influence on the control of eye movement. However, we have shown previously that these afferent signals reach various brainstem centres concerned with eye movement, notably the vestibular nuclei, and that the decerebrate pigeon is a favourable preparation in which to study their effects. If the extraocular muscle afferents do influence oculomotor control from moment-to-moment they should exert a demonstrable effect on the oculomotor nuclei. We now present evidence that extraocular muscle afferent signals do, indeed, alter the responses of units in an oculomotor nucleus (the abducens, VI nerve nucleus, which supplies the lateral rectus muscle) to horizontal, vestibular stimulation induced by sinusoidal oscillation of the bird. Such stimuli evoke a vestibulo-ocular reflex in the intact bird. The extraocular stretch receptors were activated by passive eye movement within the pigeon's saccadic range; such movements modified the vestibular responses of all 19 units studied which were all, histologically, in the abducens nucleus. The magnitude of the effects, purely inhibitory in 15 units, depended both on the amplitude and the velocity of the eye movement and most units showed selectivity for particular combinations of plane (e.g. horizontal versus vertical) and direction (e.g. rostral versus caudal) of eye movement. The results show that an afferent signal from the extraocular muscles influences vestibularly driven activity in the abducens nucleus to which it carries information related to amplitude, velocity, plane and direction of eye movement in the saccadic range. They thus strongly support the view that extraocular afferent signals are involved in the control of eye movement.     

Neuronal pathways for the lingual reflex in the Japanese toad

1. Anuran tongue is controlled by visual stimuli for releasing the prey-catching behavior ('snapping') and also by the intra-oral stimuli for eliciting the lingual reflex. To elucidate the neural mechanisms controlling tongue movements, we analyzed the neuronal pathways from the glossopharyngeal (IX) afferents to the hypoglossal (XII) tongue-muscle motoneurons. 2. Field potentials were recorded from the bulbar dorsal surface over the fasciculus solitarius (fsol) to the electrical stimulation of the ipsilateral IX nerve. They were composed of three successive negative waves: S1, S2 and N wave. The S1 and S2 waves followed successive stimuli applied at short intervals (10 ms or less), whereas the N wave was strongly suppressed at intervals shorter than 500 ms. Furthermore, the S1 wave had lower threshold than the S2 wave. 3. Orthodromic action potentials were intra-axonally recorded from IX afferent fibers in the fsol to the ipsilateral IX nerve stimuli. Two peaks found in the latency distribution histogram of these action potentials well coincided with the negative peaks of the S1 and the S2 waves of the simultaneously recorded field potentials. Therefore, the S1 and S2 waves should represent the compound action potentials of two groups of the IX afferent fibers with different conduction velocities. 4. Ipsilateral IX nerve stimuli elicited excitatory postsynaptic potentials (EPSPs) in the tongue-protractor motoneurons (PMNs) and the tongue-retractor motoneurons (RMNs). Inhibitory postsynaptic potentials were not observed. 5. The EPSPs recorded in PMNs had mean onset latencies of 6.4 ms measured from the negative peaks of the S1 wave. The EPSPs were facilitated when paired submaximal stimuli were applied at intervals shorter than 20 ms, but were suppressed at intervals longer than 30 ms. Furthermore, the EPSPs were spatially facilitated when peripherally split two bundles of the IX nerve were simultaneously stimulated. 6. On the other hand, the EPSPs recorded in RMNs had shorter onset latencies, averaging 2.5 ms. In 14 of 43 RMNs, early and late EPSP components could be reliably discriminated. The thresholds for the early EPSP components were as low as those for the S1 waves, whereas for the late EPSP components the thresholds were usually higher than those for the S2 waves.(ABSTRACT TRUNCATED AT 400 WORDS)     

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.     

Changes in postsynaptic responses in spinal motoneurons during repetitive stimulation of the locus coeruleus

Experiments on cats anesthetized with chloralose showed that repetitive stimulation of the locus coeruleus is accompanied by a decrease in IPSPs evoked by stimulation of flexor reflex afferents in extensor motoneurons. The effect appeared 600 msec after the beginning of stimulation and reached its maximum after 1500–2000 msec. Repetitive stimulation of the locus coeruleus did not change the membrane potential and did not affect EPSPs or IPSPs evoked by stimulation of low-threshold muscle afferents; EPSPs due to activation of high-threshold cutaneous and muscle afferents likewise remained unchanged. Repetitive stimulation of more central regions of the brain stem was accompanied not only by a decrease in IPSPs evoked by stimulation of flexor reflex afferents in extensor motoneurons, but also by a decrease in amplitude of EPSPs arising in response to stimulation of these same afferents in flexor motoneurons. These effects were not connected with activation of monoaminergic structures, for unlike effects arising during stimulation of the locus coeruleus, they were also found in previously reserpinized animals.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 14, No. 1, pp. 51–59, January–February, 1982.     

Convergence of mechanosensory inputs onto neuromodulatory serotonergic neurons in the leech

By the frequency-dependent release of serotonin, Retzius neurons in the leech modulate diverse behavioral responses of the animal. However, little is known about how their firing pattern is produced. Here we have analyzed the effects of mechanical stimulation of the skin and intracellular stimulation of mechanosensory neurons on the electrical activity of Retzius neurons. We recorded the electrical activity of neurons in ganglia attached to their corresponding skin segment by segmental nerve roots, or in isolated ganglia. Mechanosensory stimulation of the skin induced excitatory synaptic potentials (EPSPs) and action potentials in both Retzius neurons in a ganglion. The frequency and duration of responses depended on the strength and duration of the skin stimulation. Retzius cells responded after T and P cells, but before N cells, and their sustained responses correlated with the activity of P cells. Trains of five impulses at 10 Hz in every individual T, P, or N cell in isolated ganglia produced EPSPs and action potentials in Retzius neurons. Responses to T cell stimulation appeared after the first impulse. In contrast, the responses to P or N cell stimulation appeared after two or more presynaptic impulses and facilitated afterward. The polysynaptic nature of all the synaptic inputs was shown by blocking them with a high calcium/magnesium external solution. The rise time distribution of EPSPs produced by the different mechanosensory neurons suggested that several interneurons participate in this pathway. Our results suggest that sensory stimulation provides a mechanism for regulating serotonin-mediated modulation in the leech.     

Excitation of accessory motoneurons by stimulation of Deiters' nucleus in the cat

Acute experiments on cats under chloralose-pentobarbital anesthesia showed that application of single stimuli to Deiters' nucleus evoked monosynaptic EPSPs in motoneurons of the accessory nucleus. Latent periods of EPSPs ranged from 1.3 to 2.3 msec (mean 1.8±0.3 msec), their rise time was 0.5–1.0 msec, and their duration 7–10 msec. During repetitive stimulation the EPSPs were weakly potentiated, but with an increase in the strength of stimulation applied to Deiters' nucleus they readily changed into action potentials. In some motoneurons polysynaptic EPSPs with latent periods of the order of 6.0 msec appeared on the descending phase of these EPSPs.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 13, No. 5, pp. 515–519, September–October, 1981.     

Convergence of corticospinal and rubrospinal influences on cervical motoneurons

Synaptic responses of 121 identified cervical motoneurons to stimulation of the pyramidal tract and red nucleus were investigated by intracellular recording in cats. Responses of EPSP or EPSP-IPSP type were predominant in motoneurons of distal groups of muscles and proximal flexors, while responses of IPSP type were predominant in motoneurons of the proximal extensors. The minimal effective number of stimuli for most motoneurons was 2 or 3. The mean latent period, counted from the first stimulus in the series, was 7.86 msec for EPSPs for stimulation of the pyramidal tract and 7.91 msec for stimulation of the red nucleus, while the corresponding periods for IPSPs were 8.68 and 8.75 msec. The segmental delay of 1.3–2 msec for EPSPs and IPSPs generated in certain motoneurons in response to stimulation of both structures indicates that the shortest pathway for transmission of activity from the fibers of these tracts to the motoneurons may be disynaptic. At the same time, the possible presence of an additional neuron for most inhibitory pathways cannot be ruled out. Analysis of the results also suggests the presence of a common interneuronal apparatus for both systems.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol.3, No.6, pp. 599–608, November–December, 1971.