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.
Musienko PE, Zelenin PV, Orlovsky GN, Deliagina TG. Facilitation of postural limb reflexes with epidural stimulation in spinal rabbits. J Neurophysiol 103: 1080 -1092, 2010. First published December 16, 2009 doi:10.1152/jn.00575.2009. 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.
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.
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