Biomedical subjects

P V Zelenin

Publications and source records attributed to P V Zelenin.

13 recordsLinked to original sources

PubMed2006

Responses of reticulospinal neurons in the lamprey to lateral turns.

When swimming, the lamprey maintains a definite orientation of its body in the vertical planes, in relation to the gravity vector, as the result of postural vestibular reflexes. Do the vestibular-driven mechanisms also play a role in the control of the direction of swimming in the horizontal (yaw) plane, in which the gravity cannot be used as a reference direction? In the present study, we addressed this question by recording responses to lateral turns in reticulospinal (RS) neurons mediating vestibulospinal reflexes. In intact lampreys, the activity of axons of RS neurons was recorded in the spinal cord by implanted electrodes. Vestibular stimulation was performed by periodical turns of the animal in the yaw plane (60 degrees peak to peak). It was found that the majority of responding RS neurons were activated by the contralateral turn. By removing one labyrinth, we found that yaw responses in RS neurons were driven mainly by input from the contralateral labyrinth. We suggest that these neurons, when activated by the contralateral turn, will elicit the ipsilateral turn and thus will compensate for perturbations of the rectilinear swimming caused by external factors. It is also known that unilateral eye illumination elicits a contralateral turn in the yaw plane (negative phototaxis). We found that a portion of RS neurons were activated by the contralateral eye illumination. By eliciting an ipsilateral turn, these neurons could mediate the negative phototaxis.

Action Potentials↗

PubMed2005

Impairment and recovery of postural control in rabbits with spinal cord lesions.

The aim of this study was to characterize impairment and subsequent recovery of postural control after spinal cord injuries. Experiments were carried out on rabbits with three types of lesion--a dorsal (D), lateral (L), or ventral (V) hemisection (HS) at T(12) level. The animals were maintaining equilibrium on a platform periodically tilted in the frontal plane. We assessed the postural limb/trunk configuration from video recordings and postural reflexes in the hindquarters from kinematical and electromyographic (EMG) recordings. We found that for a few days after DHS or LHS, the animals were not able to maintain the dorsal-side-up position of their hindquarters. This ability was then gradually restored, and the dynamic postural reflexes reached the prelesion value within 2-3 wk. By contrast, a VHS almost completely abolished postural reflexes, and they did not recover for > or =7 wk. The DHS, LHS, and VHS caused immediate and slowly compensated changes in the postural limb/trunk configuration as well as gradually developing changes. After DHS, both hind limbs were placed in an abnormal rostral and medial position. After LHS, the limb on the undamaged side was turned inward and occurred at the abnormal medial position; LHS also caused a gradually developing twisting of the caudal trunk. VHS caused gradually developing extension of the ankle and knee joints. These findings show that ventral spinal pathways are of crucial importance for postural control. When a part of these pathways is spared, postural reflexes can be restored rapidly, but not the postural limb/trunk configuration. Spinal and supraspinal mechanisms responsible for postural deficits and their compensation are discussed.

Animals↗

PubMed2003

Postural control in the rabbit maintaining balance on the tilting platform.

A deviation from the dorsal-side-up body posture in quadrupeds activates the mechanisms for postural corrections. Operation of these mechanisms was studied in the rabbit maintaining balance on a platform periodically tilted in the frontal plane. First, we characterized the kinematics and electromyographic (EMG) patterns of postural responses to tilts. It was found that a reaction to tilt includes an extension of the limbs on the side moving down and flexion on the opposite side. These limb movements are primarily due to a modulation of the activity of extensor muscles. Second, it was found that rabbits can effectively maintain the dorsal-side-up body posture when complex postural stimuli are applied, i.e., asynchronous tilts of the platforms supporting the anterior and posterior parts of the body. These data suggest that the nervous mechanisms controlling positions of these parts of the body can operate independently of each other. Third, we found that normally the somatosensory input plays a predominant role for the generation of postural responses. However, when the postural response appears insufficient to maintain balance, the vestibular input contributes considerably to activation of postural mechanisms. We also found that an asymmetry in the tonic vestibular input, caused by galvanic stimulation of the labyrinths, can affect the stabilized body orientation while the magnitude of postural responses to tilts remains unchanged. Fourth, we found that the mechanisms for postural corrections respond only to tilts that exceed a certain (threshold) value.

Animals↗

PubMed2003

Comparison of the motor effects of individual vestibulo- and reticulospinal neurons on dorsal and ventral myotomes in lamprey.

In the lamprey (a lower vertebrate), motor commands from the brain to the spinal cord are transmitted through the reticulospinal (RS) and vestibulospinal (VS) pathways. The axons of larger RS neurons reach the most caudal of approximately 100 spinal segments, whereas the VS pathway does not descend below the 15th segment. This study was carried out to compare functional projections of RS and VS neurons in the rostral spinal segments that the neurons innervate together. To reveal these projections, individual RS or VS neurons were stimulated, and the responses of different groups of spinal motoneurons were recorded in ventral root branches to dorsal and ventral parts of myotomes. The responses were detected using a spike-triggered averaging technique on the background of ongoing motoneuronal activity. Individual RS and VS neurons exerted uniform effects on segmental motor output within this rostral part of the spinal cord. The effects of VS neurons on different groups of motoneurons were weaker and less diverse than those of RS neurons. The results indicate that VS neurons are able to elicit a flexion of the rostral part of the body and to turn the head in different planes without affecting more caudal parts. By contrast, larger RS neurons can elicit head movement only together with movement of a considerable part of the body and thus seem to be responsible for formation of gross motor synergies.

Animals↗

PubMed2001

Heterogeneity of the population of command neurons in the lamprey.

The effects of signals transmitted from the brain to the spinal locomotor networks by a population of command neurons are determined by specific functional projections of each individual neuron. To reveal these projections, we used a simple vertebrate model, the lamprey, in which responses of the spinal networks to spikes in single reticulospinal axons were detected by using the spike-triggered averaging of the motoneuronal activity. We found that individual neurons exert a uniform effect on the segmental motor output along the whole extent of their axons. Twenty different patterns of effect, that is, combinations of influences on the segmental motoneuron pools, were found. The widespread projections and heterogeneity of the population of command neurons present a basis for formation of different gross motor synergies.

Action Potentials↗

PubMed2001

Modeling postural control in the lamprey.

A phenomenological model of the mechanism of stabilization of the body orientation during locomotion (dorsal side up) in the lamprey is presented. The mathematical modeling is based on experimental results obtained during investigations of postural control in lampreys using a combined in vivo and robotics approach. The dynamics of the model agree qualitatively with the experimental data. It is shown by computer simulations that postural correction commands from reticulospinal neurons provide information sufficient to stabilize body orientation in the lamprey. The model is based on differences between the effects exerted by the vestibular apparatus on the left and the right side.

Animals↗

PubMed2000

Projection pattern and target selection of Clione limacina motoneurons sprouting within an intact environment.

In the pteropod mollusc Clione limacina, two groups of locomotor motoneurons, located in the pedal ganglion, innervate the dorsal and ventral muscle layers of the ipsilateral wing through the wing nerve. Separate branches of this nerve go either only to the dorsal muscle layer or only to the ventral one. In the present study, growth of novel neurites of the wing motoneurons was induced by cutting the wing nerve. In addition, all other peripheral nerves and connectives of the pedal ganglion were cut, except for the pedal commissure to the contralateral pedal ganglion. Thus, the neurites were allowed to grow only towards the contralateral pedal ganglion. We have found that the novel neurites, entering the contralateral pedal ganglion, were capable of growing everywhere inside the central nervous system (CNS) and into any peripheral nerve. However, they preferred the wing nerve. This finding suggests that the preference is caused by the guiding cues in the wing nerve or the attractive influence of the wing muscles. Because the contralateral pedal ganglion and nerves were left intact, the growth direction of the new neurites could be determined only by factors permanently existing in the CNS, rather than induced by nerve injury or muscle denervation. Within the wing nerve, the neurites could not discriminate between the nerve branches going to the dorsal and ventral muscle layers. They formed synapses on muscles of both layers, despite the fact that the muscles were innervated by their own motoneurons.

Animals↗

PubMed2000

Crossed reciprocal inhibition evoked by electrical stimulation of the lamprey spinal cord.

Activation of a motoneuron pool is often accompanied by inhibition of the antagonistic pool through a system of reciprocal inhibition between the two parts of the neuronal network controlling the antagonistic pools. In the present study, we describe the activity of such a system in the isolated spinal cord of the lamprey, when a tonic motor output is evoked by extracellular stimulation (0.5-1 s train of pulses, 20 Hz) of either end of the spinal cord. With two electrodes symmetrically positioned in relation to the midline, stimulation with either of them separately elicited prolonged (1-5 s) ipsilateral ventral root activity. Activity could be abolished by stronger, simultaneously applied, stimulation of the contralateral side of the cord, suggesting that reciprocal inhibition between hemisegments operates when a tonic motor output is generated. Simultaneous stimulation of both sides of the spinal cord with a single electrode with a large tip (300-400 microm in diameter), positioned over the anatomical midline, elicited inconsistent right-side, leftside, or bilateral ventral root responses. A minor displacement (10-20 microm) to the left or right from the midline resulted in activation of ipsilateral motoneurons, whereas the contralateral motoneurons were silent. These findings indicate that a small asymmetry in the excitatory drive to the left and right spinal hemisegments can be further amplified by reciprocal inhibition between the hemisegments. Longitudinal splitting of the spinal cord along the midline resulted in reduced reciprocal inhibition between the hemisegments separated by the lesion. The reduction was proportional to the extent of the split. The inhibition was abolished when the split reached nine segments in length. From these experiments, the longitudinal distribution of the commissural axons responsible for inhibition of contralateral motor output could be estimated.

Animals↗

PubMed2000

Activity of reticulospinal neurons during locomotion in the freely behaving lamprey.

The reticulospinal (RS) system is the main descending system transmitting commands from the brain to the spinal cord in the lamprey. It is responsible for initiation of locomotion, steering, and equilibrium control. In the present study, we characterize the commands that are sent by the brain to the spinal cord in intact animals via the reticulospinal pathways during locomotion. We have developed a method for recording the activity of larger RS axons in the spinal cord in freely behaving lampreys by means of chronically implanted macroelectrodes. In this paper, the mass activity in the right and left RS pathways is described and the correlations of this activity with different aspects of locomotion are discussed. In quiescent animals, the RS neurons had a low level of activity. A mild activation of RS neurons occurred in response to different sensory stimuli. Unilateral eye illumination evoked activation of the ipsilateral RS neurons. Unilateral illumination of the tail dermal photoreceptors evoked bilateral activation of RS neurons. Water vibration also evoked bilateral activation of RS neurons. Roll tilt evoked activation of the contralateral RS neurons. With longer or more intense sensory stimulation of any modality and laterality, a sharp, massive bilateral activation of the RS system occurred, and the animal started to swim. This high activity of RS neurons and swimming could last for many seconds after termination of the stimulus. There was a positive correlation between the level of activity of RS system and the intensity of locomotion. An asymmetry in the mass activity on the left and right sides occurred during lateral turns with a 30% prevalence (on average) for the ipsilateral side. Rhythmic modulation of the activity in RS pathways, related to the locomotor cycle, often was observed, with its peak coinciding with the electromyographic (EMG) burst in the ipsilateral rostral myotomes. The pattern of vestibular response of RS neurons observed in the quiescent state, that is, activation with contralateral roll tilt, was preserved during locomotion. In addition, an inhibition of their activity with ipsilateral tilt was clearly seen. In the cases when the activity of individual neurons could be traced during swimming, it was found that rhythmic modulation of their firing rate was superimposed on their tonic firing or on their vestibular responses. In conclusion, different aspects of locomotor activity-initiation and termination, vigor of locomotion, steering and equilibrium control-are well reflected in the mass activity of the larger RS neurons.

Animals↗

PubMed2000

Postural control in the lamprey: A study with a neuro-mechanical model.

The swimming lamprey normally maintains the dorsal-side-up orientation due to activity of the postural control system driven by vestibular organs. Commands for postural corrections are transmitted from the brain stem to the spinal cord mainly by the reticulospinal (RS) pathways. As shown in previous studies, RS neurons are activated by contralateral roll tilt, they exhibit a strong dynamic response, but much weaker static response. Here we test a hypothesis that decoding of these commands in the spinal cord is based on the subtraction of signals in the left and right RS pathways. In this study, we used a neuro-mechanical model. An intact lamprey was mounted on a platform that restrained its postural activity but allowed lateral locomotor undulations to occur. The activity in the left and right RS pathways was recorded by implanted electrodes. These natural biological signals were then used to control an electrical motor rotating the animal around its longitudinal axis toward the stronger signal. It was found that this "hybrid" system automatically stabilized a normal orientation of the lamprey in the gravitational field. The system compensated for large postural disturbances (lateral tilt up to +/-180 degrees ) due to wide angular zones of the gravitational sensitivity of RS neurons. In the nonswimming lamprey, activity of RS neurons and their vestibular responses were considerably reduced, and the system was not able to stabilize the normal orientation. However, the balance could be restored by imposing small oscillations on the lamprey, which elicited additional activation of the vestibular organs. This finding indicates that head oscillations caused by locomotor movements may contribute to postural stabilization. In addition to postural stabilization, the neuro-mechanical model reproduced a number of postural effects characteristic of the lamprey: 1) unilateral eye illumination elicited a lateral tilt ("dorsal light response") due to a shift of the equilibrium point in the vestibular-driven postural network; 2) removal of one labyrinth resulted in a loss of postural control due to an induced left-right asymmetry in the vestibulo-reticulospinal reflexes, which 3) could be compensated for by asymmetrical visual input. The main conclusion of the present study is that natural supraspinal commands for postural corrections in the roll plane can be effectively decoded on the basis of subtraction of the effects of signals delivered by the left and right RS pathways. Possible mechanisms for this transformation are discussed.

Animals↗

PubMed1999

Axotomized neurons of the pteropod mollusc Clione limacina develop novel sites of transmitter release in the absence of their normal muscle target.

Neural network for rhythmic wing movements in the swimming mollusc Clione limacina is a well-studied system. After nerve transection the efferent wing neurons cannot reach muscles and consequently display intensive central sprouting. In the present work it was shown that two types of efferent neurons with different neurotransmitters: acethylcholinergic locomotor motoneurons and serotonergic modulatory efferent neurons when deprived of their normal targets, release their neurotransmitter intended for peripheral muscles, in the unusual compartment--neuropile. Such 'unauthorized' release of neurotransmitter may cause nervous system dysfunctions in the damaged brain of other animals.

Acetylcholine↗

PubMed1999

Selective regeneration of the neuromuscular connections in the pteropod mollusc Clione limacina.

In the pteropod mollusc Clione limacina, two different groups of motoneurons innervate two physiologically identical wing muscles (dorsal and ventral). When motoneuron axons are crushed in the nerve of whole animals regeneration starts. In its course motoneurons initially project to both the correct and incorrect muscles. Then incorrect connections and neurites are eliminated and the original innervation is restored. Here we investigated neuromuscular regeneration when one of the muscles was removed or the muscle size was reduced. Motoneurons formed both correct and incorrect connections not only in vitro when the pedal ganglion was attached to only one of two wing muscles, but also in whole-animal 'no choice' experiments when only one muscle was available for reinnervation. In these experiments incorrect connections were stable and were not eliminated at the later stages, as happened in experiments in which both muscles were accessible. In whole-animal experiments with reduced size of the muscles, a normal pattern of regeneration was conserved although not all incorrect connections were eliminated. Thus, in the course of regeneration: (i) locomotor motoneurons make connections with both correct and incorrect muscles; (ii) if for some group of motoneurons the correct targets are unavailable, the incorrect connections survive and become stable; (iii) if both groups of motoneurons have a choice between the correct and incorrect targets, initial mixed innervation is replaced by purely correct innervation; (iv) elimination of incorrect synapses could be a result of the competition between correct and incorrect synapses of the same neuron.

Action Potentials↗

PubMed1995

Formation of connections between cultured identified neurones from the pleural ganglion of the pteropod mollusc Clione limacina.

A cluster of electrically interconnected neurosecretory cells (the 'white cells') involved in the control of reproductive behavior was identified in the right pleural ganglion of the marine mollusc, Clione limacina. Pleural ganglia also contain large neurons (PL1 and PL2) having no connections with each other and with the white cells. Most isolated white cells put into the simple unconditioned medium (50% L-15) adhered to the bottom of uncoated dishes and demonstrated neurite outgrowth for 7-10 days. If growing processes overlapped, the white cells formed electrical connections with each other, but they formed no connections with the PL1 and PL2 neurons. It is concluded that in the case which was under study cellular intrinsic properties were sufficient for the formation of 'correct' connections between neurones.

Animals↗