Increase of extensor tonus of forelimbs in chronic cats with bilateral serial hemisections of the spinal cord at different levels

An increase of extensor tonus of the forelimbs was observed in chronic cats with serial double hemisections of the spinal cord, first at a lower thoracic level followed by a contralateral hemisection at the midthoracic cord at intervals of 5-18 weeks. A similar increase in forelimb extensor tonus was observed in chronic cats with serial double hemisections, first at the high cervical cord followed by a contralateral hemisection of the mid-thoracic cord at intervals of 3-7 weeks. The results suggest that the augmentation of extensor tonus was brought about by release of bulbar centers from ascending inhibitory mechanisms from the lower spinal cord.     

Control of fine movements mediated by propriospinal neurons

Studies of the role of the C3/C4 propriospinal system of spinal cord interneurons in the control of fine movements and the processes compensating motor deficiency after lesioning of the cortico- and rubrospinal tracts at the level of segments C2 and C5 were studied in cats. These experiments showed that after lesioning of the cortico- and rubrospinal tracts at the level of C5, the C3/C4 propriospinal system played a key role in recovery processes, while after lesioning at the level of C2, the leading role was played by ipsilateral tracts in the ventral part of the spinal cord. In addition, the propriospinal system demonstrated a significant level of plasticity and was able to provide complete control of a number of fine voluntary movements.Translated from Rossiiskii Fiziologicheskii Zhurnal imeni I. M. Sechenova, Vol. 89, No. 9, pp. 1058–1066, September, 2003.     

Propriospinal neurons are sufficient for bulbospinal transmission of the locomotor command signal in the neonatal rat spinal cord

We recently showed that propriospinal neurons contribute to bulbospinal activation of locomotor networks in the in vitro neonatal rat brainstem–spinal cord preparation. In the present study, we examined whether propriospinal neurons alone, in the absence of long direct bulbospinal transmission to the lumbar cord, can successfully mediate brainstem activation of the locomotor network. In the presence of staggered bilateral spinal cord hemisections, the brainstem was stimulated electrically while recording from lumbar ventral roots. The rostral hemisection was located between C1 and T3 and the contralateral caudal hemisection was located between T5 and mid-L1. Locomotor-like activity was evoked in 27% of the preparations, which included experiments with staggered hemisections placed only two segments apart. There was no relation between the likelihood of developing locomotor-like activity and the distance separating the two hemisections or specific level of the hemisections. In some experiments, where brainstem stimulation alone was ineffective, neurochemical excitation of propriospinal neurons (using 5-HT and NMDA) at concentrations subthreshold for producing locomotor-like activity, promoted locomotor-like activity in conjunction with brainstem stimulation. In other experiments, involving neither brainstem stimulation nor cord hemisections, the excitability of propriospinal neurons in the cervical and/or thoracic region was selectively enhanced by bath application of 5-HT and NMDA or elevation of bath K⁺ concentration. These manipulations produced locomotor-like activity in the lumbar region. In total, the results suggest that propriospinal neurons are sufficient for transmission of descending locomotor command signals. This observation has implications for regeneration strategies aimed at restoration of locomotor function after spinal cord injury.     

Characteristics of H- and M-waves recorded from rat forelimbs

The Hoffman reflex (H-reflex) is a useful tool for studying the functional aspects of the spinal cord without anesthesia and/or damage to the body. H-reflex studies are performed mainly in the hindlimbs. The purpose of the present study was to evaluate the characteristics of the H-reflex in the forelimbs and hindlimbs in rats anesthetized with ketamine–HCl. H- and M-waves were recorded from the interosseous muscles after electrical stimulation of the n. lateral plantar of the hindlimb and n. medialis of the forelimb. Hmax/Mmax values were significantly smaller in the forelimbs than in the hindlimbs. Furthermore, paired-pulse attenuation tended to be stronger in the forelimbs than in the hindlimbs. These findings suggest that control by descending and/or propriospinal pathways is stronger in the forelimbs than in the hindlimbs in rats.     

Different patterns of fore-hindlimb coordination during overground locomotion in cats with ventral and lateral spinal lesions

The effect of large, low thoracic (T10–T11), partial spinal lesions involving the ventral quadrants of the spinal cord and, to a different extent, the dorsolateral funiculi, on fore-hindlimb coordination was examined in cats walking overground at moderate speeds (40–100 cm/s). Three different forms of impairment of fore-hindlimb coordination depending on the extent of the lesions, were observed. Lesions sparing the dorsolateral or the ventral funiculus on one side preserved the equality of the fore- and hindlimb locomotor rhythms but changed the coupling between the movements of both girdles as compared to intact animals. Larger lesions in which, in addition to the ventral quadrants of the spinal cord, also major parts of the dorsolateral funiculi were destroyed elicited episodes of rhythm oscillations in both girdles, which appeared at the background of a small difference in these rhythms. Lesions destroying almost the whole spinal cord induced a permanent difference (about 200 ms) in the step cycle duration of the fore- and the hindlimbs. However, even in these animals some remnant form of fore-hindlimb coordination was found. The results suggest that dorsolateral funiculi play a major role in preserving the equality of rhythms in the foreand the hindlimbs, while lesions of the ventral quadrants change the coupling between limbs.     

Neurochemical excitation of propriospinal neurons facilitates locomotor command signal transmission in the lesioned spinal cord

Previous studies of the in vitro neonatal rat brain stem-spinal cord showed that propriospinal relays contribute to descending transmission of a supraspinal command signal that is capable of activating locomotion. Using the same preparation, the present series examines whether enhanced excitation of thoracic propriospinal neurons facilitates propagation of the locomotor command signal in the lesioned spinal cord. First, we identified neurotransmitters contributing to normal endogenous propriospinal transmission of the locomotor command signal by testing the effect of receptor antagonists applied to cervicothoracic segments during brain stem-induced locomotor-like activity. Spinal cords were either intact or contained staggered bilateral hemisections located at right T1/T2 and left T10/T11 junctions designed to abolish direct long-projecting bulbospinal axons. Serotonergic, noradrenergic, dopaminergic, and glutamatergic, but not cholinergic, receptor antagonists blocked locomotor-like activity. Approximately 73% of preparations with staggered bilateral hemisections failed to generate locomotor-like activity in response to electrical stimulation of the brain stem alone; such preparations were used to test the effect of neuroactive substances applied to thoracic segments (bath barriers placed at T3 and T9) during brain stem stimulation. The percentage of preparations developing locomotor-like activity was as follows: 5-HT (43%), 5-HT/N-methyl-D-aspartate (NMDA; 33%), quipazine (42%), 8-hydroxy-2-(di-n-propylamino)tetralin (20%), methoxamine (45%), and elevated bath K(+) concentration (29%). Combined norepinephrine and dopamine increased the success rate (67%) compared with the use of either agent alone (4 and 7%, respectively). NMDA, Mg(2+) ion removal, clonidine, and acetylcholine were ineffective. The results provide proof of principle that artificial excitation of thoracic propriospinal neurons can improve supraspinal control over hindlimb locomotor networks in the lesioned spinal cord.     

Some properties of sympathetic neuron inhibition by depressor area and intraspinal stimulation

In Nembutal anesthesized cats single shock stimulation of the depressor area of the medulla oblongata evoked inhibition of spontaneous and glutamate-evoked activity of sympathetic preganglionic units. Single shocks to the lateral funiculus of the cervical or upper thoracic spinal cord in acute spinal cats evoked inhibition of the spontaneous and glutamate-evoked activity of single units and of the segmental reflex mass discharge evoked by spinal afferent stimulation. Cats studied 4 to 6 weeks after a complete transection of the spinal cord also showed, on stimulation of the lateral funiculus below the transection, an inhibition of the segmental reflex with time course similar to that seen in the acute spinal state, but of lower threshold and greater intensity. These results suggest that the inhibitory coupling between supraspinal levels and sympathetic preganglionic units is mediated, at least in part, by propriospinal neuronal system which survive after chronic spinal section. On the assumption that the observed changes in the properties of inhibition are due to plastic changes consequent to partial denervation the results also suggest that continuous descending tracts exist, and that both the continuous and the propriospinal descending tracts may be converging onto some common neural element.     

Control of Locomotor Activity in Humans and Animals in the Absence of Supraspinal Influences

Electrical epidural stimulation of the dorsal surface of the spinal cord at the level of the second lumbar segment induced step-like movements accompanied by the corresponding electromyographic activity in the leg muscles in patients lacking supraspinal influences as a result of vertebral trauma. Triggering of stepping movements was shown to occur with particular stimulation parameters. The results provide evidence that in humans, as in other mammals, the spinal cord contains a network of interneurons acting as generators of stepping movements and producing coordinated patterns of movement activity. Experiments on chronic spinal cats demonstrated the leading role of the propriospinal system of the spinal cord in activating the spinal generators of stepping in response to epidural stimuli.     

Initiation of Locomotor Activity in Spinal Cats by Epidural Stimulation of the Spinal Cord

Acute and chronic experiments on lower spinal (T10–T12) cats were performed to investigate the effects of epidural stimulation of the dorsal surface of the spinal cord on the initiation of locomotor activity. A zone located at the border between segments L4 and L5 was identified, stimulation of which induces locomotor activity. The parameters of epidural stimulation of the spinal cord effective in activating the stepping movement generator were identified. Epidural stimulation leading to the initiation of movement activity was shown to depend on intracentral and peripheral mechanisms activating the segmental, intersegmental and propriospinal reflex systems of the spinal cord. A leading role was demonstrated for the propriospinal system of the dorsolateral funiculi in activating the generators of stepping movements in epidural stimulation of the spinal cord.     

Attributes of quiet stance in the chronic spinal cat

Standing is a dynamic task that requires antigravity support of the body mass and active regulation of the position of the body center of mass. This study examined the extent to which the chronic spinal cat can maintain postural orientation during stance and adapt to changes in stance distance (fore-hindpaw separation). Intact cats adapt to changes in stance distance by maintaining a constant horizontal orientation of the trunk and changing orientation of the limbs, while keeping intralimb geometry constant and aligning the ground reaction forces closely with the limb axes. Postural adaptation was compared in four cats before and after spinalization at the T(6) level, in terms of the forces exerted by each paw against the support, body geometry (kinematics) and electromyographic (EMG) activity recorded from chronic, indwelling electrodes, as well as the computed net torques in the fore and hindlimbs. Five fore-hindpaw distances spanning the preferred distance were tested before spinalization, with a total range of 20 cm from the shortest to the longest stance. After spinalization, the cats were trained on a daily basis to stand on the force platform, and all four cats were able to support their full body weight. Three of the four cats could adapt to changes in stance distance, but the range was smaller and biased toward the shorter distances. The fourth cat could stand only at one stance distance, which was 8 cm shorter than the preferred distance before spinalization. All cats shifted their center of pressure closer to the forelimbs after spinalization, but the amount of shift could largely be accounted for by the weight loss in the hindquarters. The three cats that could adapt to changes in stance distance used a similar strategy as the intact cat by constraining the trunk and changing orientation of the limb axes in close relation with the forces exerted by each limb. However, different postures in the fore- and hindlimbs were adopted, particularly at the scapula (more extended) and pelvis (tipped more anteriorly). Other changes from control included a redistribution of net extensor torque across the joints of the forelimb and of the hindlimb. We concluded that the general form of body axis orientation is relatively conserved in the spinal cat, suggesting that the lumbosacral spinal circuitry includes rudimentary set points for hindlimb geometry. Both mechanical and neural elements can contribute toward maintaining body geometry through stiffness regulation and spinal reflexes.     

Interlimb postural coordination in the standing cat

The dorsal-side-up body posture in standing quadrupeds is maintained by coordinated activity of four limbs. We studied this coordination in the cat standing on the platform periodically tilted in the frontal plane. By suspending different body parts, we unloaded one, two, or three limbs. The activity of selected extensor muscles and the contact forces under the limbs were recorded. With all four limbs on the platform, extensors of the fore- and hindlimbs increased their activity in parallel during ipsilateral downward tilt. With two forelimbs on the platform, this muscular pattern persisted in the forelimbs and in the suspended hindlimbs. With two hindlimbs on the platform, the muscular pattern persisted only in the hindlimbs, but not in the suspended forelimbs. These results suggest that coordination between the two girdles is based primarily on the influences of the forelimbs upon the hindlimbs. However, these influences do not necessarily determine the responses to tilt in the hindlimbs. This was demonstrated by antiphase tilting of the fore- and hindquarters. Under these conditions, the extensors of the fore- and hindlimbs appeared uncoupled and modulated in antiphase, suggesting an independent control of posture in the fore- and hindquarters. With only one limb supporting the shoulder or hip girdle, a muscular pattern with normal phasing was observed in both limbs of that girdle. This finding suggests that reflex mechanisms of an individual limb generate only a part of postural corrections; another part is produced on the basis of crossed influences.     

Location of spinal cord pathways that control hindlimb movement amplitude and interlimb coordination during voluntary swimming in turtles

We performed mechanical lesions of the midbody (D2–D3; second to third postcervical spinal segments) spinal cord in otherwise intact turtles to locate spinal cord pathways that 1) activate and control the amplitude of voluntary hindlimb swimming movements and 2) coordinate hindlimb swimming with the movement of other limbs. Pre- and postlesion turtles were held by a band clamp around the carapace just beneath the water surface in a clear Plexiglas tank and videotaped from below so that kinematic measurements could be made of voluntary forward swimming with motion analysis software. Movements of the forelimbs (wrists) and hindlimbs (knees and ankles) were tracked relative to stationary reference points on the plastron to obtain bilateral measurements of hip and forelimb angles as functions of time along with foot trajectories. We measured changes in limb movement amplitude, cycle period, and interlimb phase before and after spinal lesions. Our results indicate that locomotor command signals that activate and regulate the amplitude of voluntary hindlimb swimming travel primarily in the dorsolateral funiculus (DLF) at the D2–D3 level and cross over to drive contralateral hindlimb movements. This suggests that electrical stimulation of the D3 DLF, which was previously shown to evoke swimming movements in the contralateral hindlimb of low-spinal turtles, activated the same locomotor command pathways that the animal uses during voluntary behavior. We also show that forelimb–hindlimb coordination is maintained by longitudinal spinal pathways that are largely confined to the ventrolateral funiculus (VLF) and mediate phase coupling of ipsilateral limbs, presumably by interenlargement propriospinal fibers.     

Chronically isolated lumbar half spinal cord generates locomotor activities in the ipsilateral hindlimb of the cat

The lumbar spinal cord of the cat was both hemisected (at L2 or L3) and longitudinally myelotomized in order to make half lumbar cord isolated from descending as well as contralateral impulses. The chronic cats recovered the ability to stand with their two forelimbs and a hindlimb, contralateral to the hemisection, 17.2 +/- 10.8 days after the operations. Two days later the hindlimb innervated by the isolated half lumbar cord regained walking capability. Phase relationships between the fore- and hindlimb muscles during locomotion were studied by recording EMGs from bilateral triceps brachii, vastus lateris and tibialis anterior muscles. Phase relationships between bilateral triceps brachii were 0.97 +/- 0.13 pi to 1.09 +/- 0.12 pi, indicating that the two forelimbs were stepping alternately and rhythmically. Phase relationships between bilateral vastus lateralis muscles were highly variable step by step, suggesting that the stepping of the hindlimb innervated by the isolated half lumbar cord was independently carried out, possibly evoked by peripheral receptors such as joint, muscle and cutaneous receptors.     

Recovery of locomotion after ventral and ventrolateral spinal lesions in the cat. II. Effects of noradrenergic and serotoninergic drugs

The effects of serotoninergic and noradrenergic drugs (applied intrathecally) on treadmill locomotion were evaluated in two adult cats subjected to a ventral and ventrolateral spinal lesion (T13). Despite the extensive spinal lesion, severely damaging important descending pathways such as the reticulo- and vestibulospinal tracts, both cats recovered quadrupedal voluntary locomotion. As detailed in a previous paper, the locomotor recovery occurred in three stages defined as early period, when the animal could not walk with its hindlimbs, recovery period, when progressive improvement occurred, and plateau period, when a more stable locomotor performance was observed. At this latter stage, the cats suffered from postural and locomotor deficits, such as poor lateral stability, irregular stepping of the hindlimbs, and inconsistent homolateral fore- and hindlimb coupling. The present study aimed at evaluating the potential of serotoninergic and/or noradrenergic drugs to improve the locomotor abilities in the early and late stages. Both cats were implanted chronically with an intrathecal cannula and electromyographic (EMG) electrodes, which allowed determination, under similar recording conditions, of the locomotor performance pre- and postlesion and comparisons of the effects of different drugs. EMG and kinematic analyses showed that norepinephrine (NE) injected in early and plateau periods improved the regularity of the hindlimb stepping and stabilized the interlimb coupling, permitting to maintain constant locomotion for longer periods of time. Methoxamine, the alpha1-agonist (tested only at the plateau period), had similar effects. In contrast, the alpha2-agonist, clonidine, deteriorated walking. Serotoninergic drugs, such as the neurotransmitter itself, serotonin (5HT), the precursor 5-hydroxytryptophan (5HTP), and the agonist quipazine improved the locomotion by increasing regularity of the hindlimb stepping and by increasing the step cycle duration. In contrast, the 5HT1A agonist 8-hydroxy-dipropylaminotetralin (DPAT) caused foot drag in one of the cats, resulting in frequent stumbling. Injection of combination of methoxamine and quipazine resulted in maintained, regular stepping with smooth movements and good lateral stability. Our results show that the effects of drugs can be integrated to the residual voluntary locomotion and improve some of its postural aspects. However, this work shows clearly that the effects of drugs (such as clonidine) may depend on whether or not the spinal lesion is complete. In a clinical context, this may suggest that different classes of drugs could be used in patients with different types of spinal cord injuries. Possible mechanisms underlying the effect of noradrenergic and serotoninergic drugs on the locomotion after partial spinal lesions are discussed.     

Responses of interneurons in the cat cervical cord to vestibular tilt stimulation

The responses of interneurons in the cervical spinal cord of the decerebrate cat to whole-body tilt were studied with a goal of identifying spinal elements in the production of forelimb vestibular postural reflexes. Interneurons both in the cervical enlargement and at higher levels, from which propriospinal neurons have been identified, were examined, both in animals with intact labyrinths and in animals with nonfunctional semicircular canals (canal plugged). Most cervical interneurons responding to tilt respond best to rotations in vertical planes aligned within 30 degrees of the roll plane. Two to three times as many neurons are excited by side-up roll tilt as are excited by side-down roll. In cats with intact labyrinths, most responses have dynamics proportional either to (and in phase with) the position of the animal or to a sum of position and tilt velocity. This is consistent with input from both otolith organs and semicircular canals. In animals without functioning canals, the "velocity" response is absent. In a few cells (8 out of 76), a more complex response, characterized by an increasing gain and progressive phase lag, was observed. These response dynamics characterize the forelimb reflex in canal-plugged cats and have been previously observed in vestibular neurons in such preparations.     

Initiation of Locomotion in Decerebrate Cats by Pulsed Magnetic Fields Projected onto Spinal Cord Segments

The motor effects induced by pulsed magnetic fields (PMF) projected onto the lumbar and cervical spinal cord were studied in decerebrate cats. A magnetic coil (inductor) of diameter 8 cm was positioned 1–2 cm above the surface of the spinal cord. Stimulation of the spinal cord with PMF was performed in two regimes: with single impulses with an intensity of 0.5–1 T and with continuous rhythmic stimulation at a frequency of 1 Hz and an intensity of 0.5 T. Application of single stimuli to the lumbar enlargement evoked reflex responses in the proximal and distal hindlimb muscles. Rhythmic stimulation initiated locomotor activity of the limb on a running treadmill, i.e., activated the neural locomotor network of the spinal cord (stepping movement generator). Magnetic stimulation of the lumbar enlargement evoked coordinated stepping movements of the hindlimbs only. Application of PMF to the cervical enlargement induced coordinated stepping movements of all four limbs, hindlimb movements starting before forelimb movements. After cessation of magnetic stimulation, the limbs completed several further coordinated movement cycles. This is the first report of the triggering of limb stepping movement generators with PMF in decerebrate cats. The results obtained here demonstrate that the neural locomotor networks of the spinal cord can be activated noninvasively and open new perspectives for the clinical use of PMF.     

Roll tilt reflexes after vestibulospinal tract lesions

Summary The effects of lesions of the vestibulospinal tracts on vestibular reflexes evoked by roll tilt in forelimb and neck extensors were examined in decerebrate cats. Sectioning the medial longitudinal fasciculus, which contains the medial vestibulospinal tract, had no major effect on the phase of the reflex, although some gain was usually lost at high stimulus frequencies. Spinal lesions at C2–C3, both cord hemisections and more restrictive lesions which cut the lateral vestibulospinal tract, produced two major effects on the forelimb. Background EMG activity was usually abolished in the triceps ipsilateral to the lesion, with partial loss of activity in the opposite limb. The tilt reflex response in the ipsilateral limb appeared normal, although it was usually necessary to raise the background excitability of the preparation by administering L-Dopa in order to observe the reflex. In contrast, the response in the contralateral limb showed a phase reversal of 180 deg at low stimulus frequencies, implying that the reflex in intact cats receives a crossed otolith-spinal input. Responses in the neck extensors splenius and biventer, recorded from compartments caudal to the spinal lesion, were relatively unaffected.Partially supported by NASA grant NSG-2380 and PHS grants NS02619 and RR07065Recipient of NIH Postdoctoral Fellowship NS06128Supported in part by NIH Research Service Award 7524     

Effect of naloxone on transmission in excitatory and inhibitory spinal reflex pathways.

The effect of systemic administration of naloxone on transmission in hindlimb reflex pathways was investigated in acute low spinal cats by conditioning monosynaptic reflexes. A marked enhancement of excitatory effects from cutaneous, joint, group II and III muscle afferents was observed in posterior biceps and semitendinosus motoneurones in 4 out of 6 experiments. In contrast, inhibitory synaptic effects in gastrocnemius and soleus motoneurones were not enhanced except weakly in one experiment. The effects of naloxone are different from those observed after spinal cord lesions interrupting axons of a previously described group of upper lumbar propriospinal neurones, which tonically suppress reflex transmission in the acute low spinal state. It is postulated that the suppression exerted by this group of neurones is not dependent on endogenous opioid peptides.     

Loss of propriospinal neurons after spinal contusion injury as assessed by retrograde labeling

We studied the number, location and size of long descending propriospinal tract neurons (LDPT), located in the cervical enlargement (C3–C6 spinal levels), and short thoracic propriospinal neurons (TPS), located in mid-thoracic spinal cord (T5–T7 spinal levels), 2, 6 and 16 weeks following a moderate low thoracic (T9) spinal cord contusion injury (SCI; 25 mm weight drop) and subsequent injections of fluorogold into the upper lumbosacral enlargement (L2–L4 spinal levels). Retrograde labeling showed that ∼23% of LDPT and 10% of TPS neurons were labeled 2 weeks after SCI, relative to uninjured animals. No additional significant decrease in number of labeled LDPT and TPS cells was found at the later time points examined, indicating that the maximal loss of propriospinal neurons in these two subpopulations occurs within the first 2 weeks post-SCI. The distribution of labeled cells post-moderate SCI was similar to normal in terms of their location within the gray matter. However, there was a significant change in the size (cross sectional area) of labeled neurons following injury, relative to uninjured controls, indicating a loss in the number of the largest class of propriospinal neurons. Interestingly, the number of labeled LDPT and TPS neurons was not significantly different following different injury severities. Although the rostro–caudal extent of the lesion site expanded between 2 and 16 weeks following injury, there was no significant difference in the number of propriospinal neurons that could be retrogradely labeled at these time points. Possible reasons for these findings are discussed.     

Significance of peripheral feedback in the generation of stepping movements during epidural stimulation of the spinal cord

Acute experiments on decerebrate and spinal cats were performed to study the role of the peripheral afferent input from hindlimb receptors in forming the locomotor pattern during epidural stimulation of the spinal cord. Evoked electromyographic activity in the muscles of the hindlimbs was analyzed, along with the kinematic parameters of stepping movements. Epidural stimulation (20–100 μA, 5 Hz) of segments L4–5 of the spinal cord was found to elicit well coordinated walking in the hindlimbs on a moving treadmill band. When the support conditions were changed (non-moving treadmill, unsupported position), epidural stimulation initiated walking with an unstable rhythm. This was associated with a change in the overall nature of the locomotor pattern and the internal structure of the stepping cycle. Alteration of the direction of movement of the treadmill band led to the appearance of backward walking. An increase in the speed of movement of the treadmill band increased the stepping frequency, mainly due to decreases in the extensor phase. Epidural stimulation applied 2–4 h after complete transection of the spinal cord at the T8–T9 level could elicit stepping movements, but only when the treadmill was moving. The role of peripheral feedback in generating the locomotor pattern in conditions of complete disconnection from supraspinal control increased significantly. These data show that peripheral feedback during epidural stimulation of the spinal cord can define the properties of the motor output. __________ Translated from Rossiiskii Fiziologicheskii Zhurnal imeni I. M. Sechenova, Vol. 91, No. 12, pp. 1407–1420, December, 2005.