Neural mechanisms of single corrective steps evoked in the standing rabbit.

Single steps in different directions are often used for postural corrections. However, our knowledge about the neural mechanisms underlying their generation is scarce. This study was aimed to characterize the corrective steps generated in response to disturbances of the basic body configuration caused by forward, backward or outward displacement of the hindlimb, as well as to reveal location in the CNS of the corrective step generating mechanisms. Video recording of the motor response to translation of the supporting surface under the hindlimb along with contact forces and activity of back and limb muscles was performed in freely standing intact and in fixed postmammillary rabbits. In intact rabbits, displacement of the hindlimb in any direction caused a lateral trunk movement toward the contralateral hindlimb, and then a corrective step in the direction opposite to the initial displacement. The time difference between onsets of these two events varied considerably. The EMG pattern in the supporting hindlimb was similar for all directions of corrective steps. It caused the increase in the limb stiffness. EMG pattern in the stepping limb differed in steps with different directions. In postmammillary rabbits the corrective stepping movements, as well as EMG patterns in both stepping and standing hindlimbs were similar to those observed in intact rabbits. This study demonstrates that the corrective trunk and limb movements are generated by separate mechanisms activated by sensory signals from the deviated limb. The neuronal networks generating postural corrective steps reside in the brainstem, cerebellum, and spinal cord.

Effect of acute lateral hemisection of the spinal cord on spinal neurons of postural networks.

In quadrupeds, acute lateral hemisection of the spinal cord (LHS) severely impairs postural functions, which recover over time. Postural limb reflexes (PLRs) represent a substantial component of postural corrections in intact animals. The aim of the present study was to characterize the effects of acute LHS on two populations of spinal neurons (F and E) mediating PLRs. For this purpose, in decerebrate rabbits, responses of individual neurons from L5 to stimulation causing PLRs were recorded before and during reversible LHS (caused by temporal cold block of signal transmission in lateral spinal pathways at L1), as well as after acute surgical LHS at L1. Results obtained after Sur-LHS were compared to control data obtained in our previous study. We found that acute LHS caused disappearance of PLRs on the affected side. It also changed a proportion of different types of neurons on that side. A significant decrease and increase in the proportion of F- and non-modulated neurons, respectively, was found. LHS caused a significant decrease in most parameters of activity in F-neurons located in the ventral horn on the lesioned side and in E-neurons of the dorsal horn on both sides. These changes were caused by a significant decrease in the efficacy of posture-related sensory input from the ipsilateral limb to F-neurons, and from the contralateral limb to both F- and E-neurons. These distortions in operation of postural networks underlie the impairment of postural control after acute LHS, and represent a starting point for the subsequent recovery of postural functions.

[Central Pattern Generators: Mechanisms of the Activity and Their Role in the Control of "Automatic" Movements].

Central pattern generators (CPGs) are a set of interconnected neurons capable of generating a basic pattern of motor output underlying "automatic" movements (breathing, locomotion, chewing, swallowing, and so on) in the absence of afferent signals from the executive motor apparatus. They can be divided into the constitutive CPGs active throughout the entire lifetime (respiratory CPGs) and conditional CPGs controlling episodic movements (locomotion, chewing, swallowing, and others). Since a motor output of CPGs is determined by their internal organization, the activities of the conditional CPGs are initiated by simple commands coming from higher centers. We describe the structural and functional organization of the locomotor CPGs in the marine mollusk Clione limacina, lamprey, frog embryo, and laboratory mammals (cat, mouse, and rat), CPGs controlling the respiratory and swallowing movements in mammals, and CPGs controlling discharges of the electric organ in the gymnotiform fish. It is shown that in all these cases, the generation of rhythmic motor output is based both on the endogenous (pacemaker) activity of specific groups of interneurons and on interneural interactions. These two interrelated mechanisms complement each other, ensuring the high reliability of CPG functionality. We discuss how the experience obtained in studying CPGs can be used to understand mechanisms of more complex functions of the brain, including its cognitive functions.

Effects of galvanic vestibular stimulation on postural limb reflexes and neurons of spinal postural network.

Quadrupeds maintain the dorsal side up body orientation due to the activity of the postural control system driven by limb mechanoreceptors. Binaural galvanic vestibular stimulation (GVS) causes a lateral body sway toward the anode. Previously, we have shown that this new position is actively stabilized, suggesting that GVS changes a set point in the reflex mechanisms controlling body posture. The aim of the present study was to reveal the underlying neuronal mechanisms. Experiments were performed on decerebrate rabbits. The vertebral column was rigidly fixed, whereas hindlimbs were positioned on a platform. Periodic lateral tilts of the platform caused postural limb reflexes (PLRs): activation of extensors in the loaded and flexing limb and a decrease in extensor activity in the opposite (unloaded and extending) limb. Putative spinal interneurons were recorded in segments L4-L5 during PLRs, with and without GVS. We have found that GVS enhanced PLRs on the cathode side and reduced them on the anode side. This asymmetry in PLRs can account for changes in the stabilized body orientation observed in normal rabbits subjected to continuous GVS. Responses to platform tilts (frequency modulation) were observed in 106 spinal neurons, suggesting that they can contribute to PLR generation. Two neuron groups were active in opposite phases of the tilt cycle of the ipsi-limb: F-neurons in the flexion phase, and E-neurons in the extension phase. Neurons were driven mainly by afferent input from the ipsi-limb. If one supposes that F- and E-neurons contribute, respectively, to excitation and inhibition of extensor motoneurons, one can expect that the pattern of response to GVS in F-neurons will be similar to that in extensor muscles, whereas E-neurons will have an opposite pattern. We have found that ~40% of all modulated neurons meet this condition, suggesting that they contribute to the generation of PLRs and to the GVS-caused changes in PLRs.

Use of galvanic vestibular feedback to control postural orientation in decerebrate rabbits.

In quadrupeds, the dorsal-side-up body orientation during standing is maintained due to a postural system that is driven by feedback signals coming mainly from limb mechanoreceptors. In caudally decerebrated (postmammillary) rabbits, the efficacy of this system is considerably reduced. In this paper, we report that the efficacy of postural control in these animals can be restored with galvanic vestibular stimulation (GVS) applied transcutaneously to the labyrinths. In standing intact rabbits, GVS causes a lateral body sway towards the positive electrode. We used this GVS-caused sway to counteract the lateral body sway resulting from a mechanical perturbation of posture. Experiments were performed on postmammillary rabbits that stood on the tilting platform with their hindlimbs. To make the GVS value dependent on the postural perturbation (i.e., on the lateral body sway caused by tilt of the platform), an artificial feedback loop was formed in the following ways: 1) Information about the body sway was provided by a mechanical sensor; 2) The GVS current was applied when the sway exceeded a threshold value; the polarity of the current was determined by the sway direction. This simple algorithm allowed the "hybrid" postural system to maintain the dorsal-side-up orientation of the hindquarters when the platform was tilted by ± 20°. Thus, an important postural function, i.e., securing lateral stability during standing, can be restored in decerebrate rabbits with the GVS-based artificial feedback. We suggest that such a control system can compensate for the loss of lateral stability of various etiologies, and can be used for restoration of balance control in patients with impaired postural functions.

Facilitation of postural limb reflexes in spinal rabbits by serotonergic agonist administration, epidural electrical stimulation, and postural training.

In quadrupeds, spinalization in the thoracic region severely impairs postural control in the hindquarters. The goal of this study was to improve postural functions in chronic spinal rabbits by regular application of different factors: intrathecal injection of the 5-HT(2) agonist (±)-1-(2,5-dimethoxy-4-iodophenyl)-2-aminopropane hydrochloride (DOI), epidural electrical spinal cord stimulation (EES), and specific postural training (SPT). The factors were used either alone (SPT group) or in combination (DOI+SPT, EES+SPT, and DOI+EES+SPT groups) or not used (control group). It was found that in none of these groups did normal postural corrective movements in response to lateral tilts of the supporting platform reappear within the month of treatment. In control group, reduced irregular electromyographic (EMG) responses, either correctly or incorrectly phased in relation to tilts, were observed. By contrast, in DOI+SPT and EES+SPT groups, a gradual threefold increase in the proportion of correctly phased EMG responses (compared with control) was observed. The increase was smaller in DOI+EES+SPT and SPT groups. Dissimilarly to these long-term effects, short-term effects of DOI and EES were weak or absent. In addition, gradual development of oscillatory EMG activity in the responses to tilts, characteristic for the control group, was retarded in DOI+SPT, EES+SPT, DOI+EES+SPT, and SPT groups. Thus regular application of the three tested factors and their combinations caused progressive, long-lasting plastic changes in the isolated spinal networks, resulting in the facilitation of spinal postural reflexes and in the retardation of the development of oscillatory EMG activity. The facilitated reflexes, however, were insufficient for normal postural functions.

Activity of motor cortex neurons during backward locomotion.

Forward walking (FW) and backward walking (BW) are two important forms of locomotion in quadrupeds. Participation of the motor cortex in the control of FW has been intensively studied, whereas cortical activity during BW has never been investigated. The aim of this study was to analyze locomotion-related activity of the motor cortex during BW and compare it with that during FW. For this purpose, we recorded activity of individual neurons in the cat during BW and FW. We found that the discharge frequency in almost all neurons was modulated in the rhythm of stepping during both FW and BW. However, the modulation patterns during BW and FW were different in 80% of neurons. To determine the source of modulating influences (forelimb controllers vs. hindlimb controllers), the neurons were recorded not only during quadrupedal locomotion but also during bipedal locomotion (with either forelimbs or hindlimbs walking), and their modulation patterns were compared. We found that during BW (like during FW), modulation in some neurons was determined by inputs from limb controllers of only one girdle, whereas the other neurons received inputs from both girdles. The combinations of inputs could depend on the direction of locomotion. Most often (in 51% of forelimb-related neurons and in 34% of the hindlimb-related neurons), the neurons received inputs only from their own girdle when this girdle was leading and from both girdles when this girdle was trailing. This reconfiguration of inputs suggests flexibility of the functional roles of individual cortical neurons during different forms of locomotion.

Facilitation of postural limb reflexes with epidural stimulation in spinal rabbits.

It is known that after spinalization animals lose their ability to maintain lateral stability when standing or walking. A likely reason for this is a reduction of the postural limb reflexes (PLRs) driven by stretch and load receptors of the limbs. The aim of this study was to clarify whether spinal networks contribute to the generation of PLRs. For this purpose, first, PLRs were recorded in decerebrated rabbits before and after spinalization at T12. Second, the effects of epidural electrical stimulation (EES) at L7 on the limb reflexes were studied after spinalization. To evoke PLRs, the vertebrate column of the rabbit was fixed, whereas the hindlimbs were positioned on the platform. Periodic lateral tilts of the platform caused antiphase flexion-extension limbs movements, similar to those observed in intact animals keeping balance on the tilting platform. Before spinalization, these movements evoked PLRs: augmentation of extensor EMGs and increase of contact force during limb flexion, suggesting their stabilizing postural effects. Spinalization resulted in almost complete disappearance of PLRs. After EES, however, the PLRs reappeared and persisted for up to several minutes, although their values were reduced. The post-EES effects could be magnified by intrathecal application of quipazine (5-HT agonist) at L4-L6. Results of this study suggest that the spinal cord contains the neuronal networks underlying PLRs; they can contribute to the maintenance of lateral stability in intact subjects. In acute spinal animals, these networks can be activated by EES, suggesting that they are normally activated by a tonic supraspinal drive.

Activity of pyramidal tract neurons in the cat during standing and walking on an inclined plane.

To keep balance when standing or walking on a surface inclined in the roll plane, the cat modifies its body configuration so that the functional length of its right and left limbs becomes different. The aim of the present study was to assess the motor cortex participation in the generation of this left/right asymmetry. We recorded the activity of fore- and hindlimb-related pyramidal tract neurons (PTNs) during standing and walking on a treadmill. A difference in PTN activity at two tilted positions of the treadmill (+/- 15 deg) was considered a positional response to surface inclination. During standing, 47% of PTNs exhibited a positional response, increasing their activity with either the contra-tilt (20%) or the ipsi-tilt (27%). During walking, PTNs were modulated in the rhythm of stepping, and tilts of the supporting surface evoked positional responses in the form of changes to the magnitude of modulation in 58% of PTNs. The contra-tilt increased activity in 28% of PTNs, and ipsi-tilt increased activity in 30% of PTNs. We suggest that PTNs with positional responses contribute to the modifications of limb configuration that are necessary for adaptation to the inclined surface. By comparing the responses to tilts in individual PTNs during standing and walking, four groups of PTNs were revealed: responding in both tasks (30%); responding only during standing (16%); responding only during walking (30%); responding in none of the tasks (24%). This diversity suggests that common and separate cortical mechanisms are used for postural adaptation to tilts during standing and walking.

Impairment of postural control in rabbits with extensive spinal lesions.

Our previous studies on rabbits demonstrated that the ventral spinal pathways are of primary importance for postural control in the hindquarters. After ventral hemisection, postural control did not recover, whereas after dorsal or lateral hemisection it did. The aim of this study was to examine postural capacity of rabbits after more extensive lesion (3/4 section of the spinal cord at T(12) level), that is, with only one ventral quadrant spared (VQ animals). They were tested before (control) and after lesion on the platform periodically tilted in the frontal plane. In control animals, tilts of the platform regularly elicited coordinated electromyographic (EMG) responses in the hindlimbs, which resulted in generation of postural corrections and in maintenance of balance. In VQ rabbits, the EMG responses appeared only in a part of tilt cycles, and they could be either correctly or incorrectly phased in relation to tilts. Because of a reduced value and incorrect phasing of EMG responses on both sides, this muscle activity did not cause postural corrective movements in the majority of rabbits, and the body swayed together with the platform. In these rabbits, the ability to perform postural corrections did not recover during the whole period of observation (< or =30 days). Low probability of correct EMG responses to tilts in most rabbits as well as an appearance of incorrect responses to tilts suggest that the spinal reflex chains, necessary for postural control, have not been specifically selected by a reduced supraspinal drive transmitted via a single ventral quadrant.

Maintenance of lateral stability during standing and walking in the cat.

During free behaviors animals often experience lateral forces, such as collisions with obstacles or interactions with other animals. We studied postural reactions to lateral pulses of force (pushes) in the cat during standing and walking. During standing, a push applied to the hip region caused a lateral deviation of the caudal trunk, followed by a return to the initial position. The corrective hindlimb electromyographic (EMG) pattern included an initial wave of excitation in most extensors of the hindlimb contralateral to push and inhibition of those in the ipsilateral limb. In cats walking on a treadmill with only hindlimbs, application of force also caused lateral deviation of the caudal trunk, with subsequent return to the initial position. The type of corrective movement depended on the pulse timing relative to the step cycle. If the force was applied at the end of the stance phase of one of the limbs or during its swing phase, a lateral component appeared in the swing trajectory of this limb. The corrective step was directed either inward (when the corrective limb was ipsilateral to force application) or outward (when it was contralateral). The EMG pattern in the corrective limb was characterized by considerable modification of the hip abductor and adductor activity in the perturbed step. Thus the basic mechanisms for balance control in these two forms of behavior are different. They perform a redistribution of muscle activity between symmetrical limbs (in standing) and a reconfiguration of the base of support during a corrective lateral step (in walking).

Effect of intrathecal administration of serotoninergic and noradrenergic drugs on postural performance in rabbits with spinal cord lesions.

Our previous studies have shown that extensive spinal lesions at T12 in the rabbit [ventral hemisection (VHS) or 3/4-section that spares one ventral quadrant (VQ)] severely damaged the postural system. When tested on the platform periodically tilted in the frontal plane, VHS and VQ animals typically were not able to perform postural corrective movements by their hindlimbs, although EMG responses (correctly or incorrectly phased) could be observed. We attempted to restore postural control in VHS and VQ rabbits by applying serotoninergic and noradrenergic drugs to the spinal cord below the lesion through the intrathecal cannula. It was found that serotonin and quipazine (5-HT1,2,3 agonist) did not re-establish postural corrective movements. However, when applied during a 10-day period after lesion, these drugs produced a twofold increase of the proportion of correct EMG responses to tilts. It was also found that methoxamine (alpha1 noradrenergic agonist), as well as the mixture of methoxamine and quipazine, did not re-establish postural corrective movements and did not increase the proportion of correct EMG responses. Serotonin (at later stages) and methoxamine induced periodical bursting in EMGs, suggesting activation of spinal rhythm-generating networks. Appearance of bursting seems to perturb normal operation of postural mechanisms, as suggested by methoxamine-induced abolishment of postural effects of quipazine. When applied in an intact animal, none of the tested drugs affected the value of postural corrections or evoked periodical bursting. We conclude that activation of the serotoninergic system (but not the noradrenergic one) causes selective enhancement of spinal postural reflexes during the earlier postlesion period.

Postural performance in decerebrated rabbit.

It is known that animals decerebrated at the premammillary level are capable of standing and walking without losing balance, in contrast to postmammillary ones which do not exhibit such behavior. The main goals of the present study were, first, to characterize the postural performance in premammillary rabbits, and, second, to activate the postural system in postmammillary ones by brainstem stimulation. For evaluation of postural capacity of decerebrated rabbits, motor and EMG responses to lateral tilts of the supporting platform and to lateral pushes were recorded before and after decerebration. In addition, the righting behavior (i.e., standing up from the lying position) was video recorded. We found that, in premammillary rabbits, responses to lateral tilts and pushes were similar to those observed in intact ones, but the magnitude of responses was reduced. During righting, premammillary rabbits assumed the normal position slower than intact ones. To activate the postural system in postmammillary rabbits, we stimulated electrically two brainstem structures, the mesencephalic locomotor region (MLR) and the ventral tegmental field (VTF). The MLR stimulation (prior to elicitation of locomotion) and the VTF stimulation caused an increase of the tone of hindlimb extensors, and enhanced their responses to lateral tilts and to pushes. These results indicate that the basic mechanisms for maintenance of body posture and equilibrium during standing are present in decerebrated animals. They are active in the premammillary rabbits but need to be activated in the postmammillary ones.

Influences of sensory input from the limbs on feline corticospinal neurons during postural responses.

The dorsal-side-up body posture of standing quadrupeds is maintained by coordinated activity of all limbs. Somatosensory input from the limbs evokes postural responses when the supporting surface is perturbed. The aim of this study was to reveal the contribution of sensory inputs from individual limbs to the posture-related modulation of pyramidal tract neurons (PTNs) arising in the primary motor cortex. We recorded the activity of PTNs from the limb representation of motor cortex in the cat maintaining balance on a platform periodically tilted in the frontal plane. Each PTN was recorded during standing on four limbs, and when two or three limbs were lifted from the platform and thus did not signal its displacement to motor cortex. By comparing PTN responses to tilts in different tests we found that the amplitude and the phase of the response in the majority of them were determined primarily by the sensory input from the corresponding contralateral limb. In a portion of PTNs, this input originated from afferents of the peripheral receptive field. Sensory input from the ipsilateral limb, as well as input from limbs of the other girdle made a much smaller contribution to the PTN modulation. These results show that, during postural activity, a key role of PTNs is the feedback control of the corresponding contralateral limb and, to a lesser extent, the coordination of posture within a girdle and between the two girdles.

Spinal and supraspinal postural networks.

Different species maintain a particular body orientation in space (upright in humans, dorsal-side-up in quadrupeds, fish and lamprey) due to the activity of a closed-loop postural control system. We will discuss operation of spinal and supraspinal postural networks studied in a lower vertebrate (lamprey) and in two mammals (rabbit and cat). In the lamprey, the postural control system is driven by vestibular input. The key role in the postural network belongs to the reticulospinal (RS) neurons. Due to vestibular input, deviation from the stabilized body orientation in any (roll, pitch, yaw) plane leads to generation of RS commands, which are sent to the spinal cord and cause postural correction. For each of the planes, there are two groups of RS neurons responding to rotation in the opposite directions; they cause a turn opposite to the initial one. The command transmitted by an individual RS neuron causes the motor response, which contributes to the correction of posture. In each plane, the postural system stabilizes the orientation at which the antagonistic vestibular reflexes compensate for each other. Thus, in lamprey the supraspinal networks play a crucial role in stabilization of body orientation, and the function of the spinal networks is transformation of supraspinal commands into the motor pattern of postural corrections. In terrestrial quadrupeds, the postural system stabilizing the trunk orientation in the transversal plane was analyzed. It consists of two relatively independent sub-systems stabilizing orientation of the anterior and posterior parts of the trunk. They are driven by somatosensory input from limb mechanoreceptors. Each sub-system consists of two closed-loop mechanisms - spinal and spino-supraspinal. Operation of the supraspinal networks was studied by recording the posture-related activity of corticospinal neurons. The postural capacity of spinal networks was evaluated in animals with lesions to the spinal cord. Relative contribution of spinal and supraspinal mechanisms to the stabilization of trunk orientation is discussed.

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.

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.

Activity of pyramidal tract neurons in the cat during postural corrections.

The dorsal side-up body orientation in quadrupeds is maintained by a postural control system. We investigated participation of the motor cortex in this system by recording activity of pyramidal tract neurons (PTNs) from limb representations of the motor cortex during postural corrections. The cat was standing on the platform periodically tilting in the frontal plane, and maintained equilibrium at different body configurations: with the head directed forward (symmetrically alternating loading of the left and right fore limbs), or with the head voluntary turned to the right or to the left (asymmetrical loading). We found that postural corrective responses to tilts included an increase of the contact forces and activity of limb extensors on the side moving down, and their decrease on the opposite side. The activity of PTNs was strongly modulated in relation to the tilt cycle. Phases of activity of individual PTNs were distributed over the cycle. Thus the cortical output mediated by PTNs appeared closely related to a highly automatic motor activity, the maintenance of the body posture. An asymmetrical loading of limbs, caused by head turns, resulted in the corresponding changes of motor responses to tilts. These voluntary postural modifications were also well reflected in the PTNs' activity. The activity of a part of PTNs correlated well with contact forces, in some others with the limb muscle activity; in still others no correlation with these variables was observed. This heterogeneity of the PTNs population suggests a different functional role of individual PTNs.

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.

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.